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Contents
lntroduction ......................................................................................................................................................................... vii
AS Level
1Cell str ucture ........ ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ......... ...... 1
1.1 Themicroscopeincellstudies ............................................................................................. 1
1.2 Cellsasthebasicunitsofliving organisms ................................................................................................................... 14
2Biologicalmolecul es ......................................................................................................................................................... 29
2 .1 Testingforbiologicalmolecules ................................................................................................................................... 29
2 .2 Carbohydratesand lipids ............................................................................................................................................... 30
2 .3 Proteinsandwater ........................................................................................................................................................ 44
3 Enzymes ............................................................................................................................................................................. 56
3 .1 Modeofactionofenzymes ........................................................................................................................................... 56
3 .2 Factorsthataffectenzymeaction ................................................................................................................................. 63
4Cell membranes and transport ••..•.•..••..•••..••..•.•..••..•••..••..•.•..••..•••..••..•.•..••..•••..••..•.•..••..•••..••..•.•..••..•••..••..•.•..••..•••..••..•.•..•• 74
4.1 Fluid mosaic membranes ................................................................................................. 74
4.2 Movementofsubstancesintoand out of cells ........................................................................... 78
5 The mitoti c ce ll cycl e.... .. .. .. ... ......... .. .. .. ... ......... .. .. .. ... ......... .. .. .. ... ......... .. .. .. ... ......... .. .. .. ... ......... .. .. .. ... ......... .. .. .. ... ......... .... 97
5 .1 Replication and division of nuclei and cells .................................................................................................................. 97
5 .2 Chromosome behaviour in mitosis .............................................................................................................................. 105
6 Nucleicacids andprotein synthesis ................................................................................................................................ 11 0
6 .1 StructureandreplicationofDNA ................................................................................................................................ 110
6 .2 Protein synthesis ......................................................................................................................................................... 118
JTransportinplants ... ..................................................................................................... 128
7 .1 5tructureoftransporttissues ...................................................................................................................................... 128
7 .2 Transportmechanisms ................................................................................................................................................. 135
8 Transport in mammals ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ..... 151
8 .1 Thecirculatorysystem ................................................................................................................................................. 151
8 .2 Theheart ...................................................................................................................................................................... 163
9 Gasexchangeand smokin g ............................................................................................................................................ 172
10 Infecti ous disease ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. ....... .. 192
10.1 lnfectiousdiseases .................................................................................................................................................... 192
10.2 Antibiotics ................................................................................................................................................................. 210
11 lmmunlty ....................................................................................................................................................................... 218
Contents
A Level
12 Energyandrespiration. ... . .... 234
12. 1 Energy ....................................................................................................................................................................... 234
12.2 Respiration ................................................................................................................................................................ 242
13 Photosynthesis ... .. .. .. ....... ..... .. .. .. ....... ..... .. .. .. ....... ..... .. .. .. ....... ..... .. .. .. ....... ..... .. .. .. ....... ..... .. .. .. ....... ..... .. .. .. ....... ..... .. .. .. ..... 261
13.1 Photosynthesisasanenergytransferprocess .......................................................................................................... 261
13.2 lnvestigationoflimitingfactors ............................................................................................................................... 273
13.3 Adaptationsforphotosynthesis ................................................................................................................................ 279
16 Inherited change .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ............ .. .. .. ..... 342
16 .1 Passageofinformationfromparenttooffspring .................................................................................................... 342
16.2 The rolesofgenesindeterminingthephenotype .................................................................................................... 350
16 .3 Genecontrol .............................................................................................................................................................. 372
17 Selecti on andevolution .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. ... .. 377
17.1 Variation .................................................................................................................................................................... 377
17.2 Natural and artificial selection .................................................................................................................................. 383
17.3 Evolution ................................................................................................................................................................... 399
Student's CD contents
Appendix 1: Background chemistry for biologists
Appendix 2: Investigations.data handling and statistics
Appendix 3: Preparing for your exam
Also,fore3chropk
• AnimemcriveteSI
• Alisiofkeyterms
• A topic summary
• Additional work on dam handling and pmcrirnl skills
• Suggested websites and funher reading
• A revisionchecklist
• Answers to all the examination-style questions
l11is page intentionally left blank
Introduction
Cambn'dge /men,atio"al AS a>1d A le,,el Biology is an excellent introduction to the subject
and a sound fou n,fation for studies beyond A Level, in funher and higher education, for
professional courses and for productive employment in the future. Successful study of this
programmegivesHfelongskiJJs,including:
• confidence in a technological world and infonned interest in scientific matters
• understanding of how sdemific theories and methods have developed
• awareness of the applications of biology in everyday life
• abilityrocommunicateeffectivcly
• concern for accuracy and precision
• awarenessoftheimportanceofobjectivity,integrity,enquiry,initiativeand inventivenes.s
• understanding of the usefulness and limitations of scientific methods and their
applications
• appreciation thm biology is affected by social, economic, technological, ethical and
To p ics 1 10 11 cover Sections 1 to 1 l of the AS Level syllabus and are for all students. AS
studentsareassessedonlyonthese.
Topics 12 to 19 cover Sections 12 to 19, the additional sections of the sylbbus for A Level
students only.
• Questions are included to assist comprehension and recall. Answers to these are given
at the back of the book. At the end of each topic. examination-style questions are given.
Answers to these are given on the CD
vii
Introduction
4' '.;:'~;] :: 1: ~ ·::;s;,!/:;Jife and all organisms are composed of one or more cells. There are two
fundamentaltypesofcell:prokaryoticandeukaryotic.
4' ~=::'::i~ : '/ ::1:~n:re intertwined and cannot be studied in isolation: observation and
enquiry, =perimentation and fieldwork are fundamental ro biology.
Author's acknowledgements
Jamindebtedtotheexperiencedintemationalreachersandthesrudentswholhavebecn
privileged to meet in Asia and in the UK in the process of preparing this material. I am especially
indebted to Christine Lea, an experienced teacher and examiner of Biology who has guided me
topic by topic on the special needs of the students for whom this book is designed.
Finally, Jamindebredtothepublishingte-amofprojecteditor,LydiaYoung,editorJoanna
Silman and designer Melissa Brunelli at Hodder Education, and to freelance editor Penny Nicholson
whose skiJJ and patience have brought together text and illustration as I have wished. I am most
graceful to them.
DrChrisdegg
Salisbury,Wiltshire,UK
June,2014
viii
AS Level
1 Cell structure
All org anisms are composed of cells. Knowle dge of the ir str uc tu re Th e use o f light microscopes is a funda me ntal skill th at is deve loped
a ndfun ctionunde rpin s much o fbiolo gy.Th e fund amenta l inth is topica nd a pplie dthroughout severa lothe r sectionsoft he
diffe re nces between e uka ryoti c a nd prokuyotic cells are ex plored sylla bus. Throughout the course, photomicrogra phs a nd electron
a nd provide useful biolog ica l background for th e topic o n Infectious m·a ogra phs from tra nsm· ss n a nd scann·ng e lect ron m· rosc:opes
di sease.Virusesare introd uced as non-cellula r structures,which sho uld be studi ed.
g ivesca ndida tes the opportunitytoc onside rwh ethe rce lls are a
fund a me nta l prope rty of life.
An understanding of the By the end of this section you should be able to:
principles of microscopy
shows why light and a) compare t he structure of typical a nima l and plant ce ll, by ma king te mporary pre parations of
electron microscopes have livemate rialandu,ingphotomkrographs
been essential in improving b) ca lculate the line ar magnificat ions of drawings, photomicrograph, and e lectron micrograph•
our kna.vledge of cells c) use an ~pi ece graticule a nd stage microme te r scale to me uu re ce ll, and be fam iliar with units
(m illimet re, microme tre, nanome t re) used in ce ll studies
d) expl ainanddistinguishbetwff nresolutionandmagnifica tion, wit hre fue nce tolight
microscopy and e lectron mkrosc:opy
e) calculate actual sizes of specime ns from drawings, photomicrograph, and e lectron micrograph,
Introducing cells
The cell is the basic unit of living matter - the smallest pan of an organism which we can say is
alive. hiscellsthatcarryouttheessentialprocessesoflife. Wethinkofthemasself-containedunits
o f strucrure and funaion. Some organisms are made of a single cell and are known as unicdlula r .
Examples of unicellu lar organisms are introduced in Figure 1.1. Jn fact. there are vast numbers of
different unicellular organisms in the living world, many with a very long evo lutionary history.
Other organisms are made o f many cells and are kno wn as multicellular organisms. Examples
o f multicellu lar organisms are the mammals and llowering plants. Much of the biology in this
1 Stat etheessential book is about multicellular organisms, including humans. and the processes that go on in these
processes characterist ic
organisms. But remember. single -celled organisms carry o ut all the essential func1ions of life too,
of living things
onlytheseocrurwithinthesinglecell.
D cellstructure
Amoeb.1 - ap<0tol(),l(loffreshwa1e,llabi!ats
,,,,.-
·-"· ''··::.,..
~:= ,rembraoe
"""'"'
Figure1.11ntroducingunicPllularOf!Janisatioo
'Plasmid,areillustratedinF,gure1.24(page24)andinfig ure19.5 (page4 59)
Cell size
Cells are extremely small - most are only visible as distinct strucrures when we llSf" a mlcr01Scope
(although a few types of cells are just large enough to be seen b)-· the naked eye).
Observation, of cells were first reported over 300 years ago, following the early development
of microscopes (see Figure 1.2). Today we use a com pound lig ht m icroscope to investigate
cell structure - perhaps you arealreadyfamiliarwiththelightmicroscopeasapieceoflaboratory
equipment. You may have used one to view living cells, such as the single-celled animal. Amoeba.
shown in Figure 1.1
Table 1.1 Units of length used inmicltl5(opy Since cells are so small, we need suitable units to measure them. The m e lre (symbol m )
isthestandardunitoflengthusedinscience. Thisisanintemationallyagreedunit,or
1 metre(m )•1000ml limetres(mm) Slunk Look at Table 1.1 below. This shows the sulxlivisions o f the metrethatweuseto
1---'~==~== (µm~(~l•~m='-1 measure cells and their contents. These units are listed in descending order of size. You
1 µm • 1000nanometres{nm) will see that each sulxlivision is ~ of the unit above it. The smallest units are probably
1 0
quite new to you ; they may take some getting used to.
The d imensions of cells are expressed in the unit called a m icrorneter or micron (µm). Notice this
unit is one thousandth (10--l) of a millimetre. This gives us a clear idea about how small cells are
2 Calrulate
when compared to the millimetre, which you can see on a standard ruler.
a howmanycellsof
Bacteria are really small, typically 0.5--lOµm in size, whereas the cells of plams and animals are
lOOµmdiameter1\ill
fitsidebysidealong often in the range SO-lSOµm, or larger. Jn fact, the lengths of the unicellular organisms shown in
a millimetre Figurel . l are approximately:
b themagnification Cbk"")'d amo>1as 30µm
of the image of
400 µm(butitsshapeandthereforelengthvariesgreatly)
Escherichiacoliin
Figurel.1
1.1The microscope in cell studies I
Robert Hook•(1662),anexpeflmechal!kaodooeo!
1r.etoondersoitheRoyalSocietyinloodoo,was
lascinatedbymicmsropy.rH'devisedarnmpoor>d
microscope,aodused~toobservethestrnctu,eofarl
:!'."'~~as~f=~~c:, 1~~~l,"."'"'ured
LNuw•nho•k's microscop•
~~Z!'::'~~~~~~~~~i!"~~~d
<aooom-=tsoltirtjparlide<(polleng,ains,in
:.::::.=::·:::::~,,. ~~
Ge,manbiologists,estal>lishedcel!sasthenatu,alunrt
o<<om~oruoom,o<'-"'"'™"·c-,,,,o,.,o=. ~
8
aodenli,eanina!saodplantsa<eagg<egatesoithese
organism,arrangedtodefinitelaws'
Rudolf Virchcw(1856taGermar1Jl,llhokxjst.est.lblished
1r.eideathat cellsarise(}ff'jbydivisionofexi,1ir,gc,..;I:,
Cell theory
Many biologists helped to develop the idea that living things are made of cells. This idea has
become known as the cell theory. This concepi evolved gradually during the nineteenth century,
following a steadily accelerating pace in the development d m k r oKop y and b iochemistry. You
can see a summary of these developments in Figure 1.2.
Today we recognise that the statement that cells are the unit of structure and function in living
thingsre3llycontainsthreebasicidea.s.
• CellsarethebuUdingblocksofstructure in living things.
laf~l"'"""""-"'IVacoole,
surroonded byamembrane
Figure1.3Plantand~nimalceHsfmmmulticellularOf!jilnisms
D cellstructure
Microscopy
A microscope is used to prcxluce a magnified image of an object or specimen. Tcxlay. cells can be
observed by two fundamentally differem types of microscopy:
• the compound light microscope, using visible light;
• theelectronmicroscope.usingabeamofelectrons.
In this course you will be using the light microscope frequently, and we start here, using it to
observe temporary preparations of living cells. later, we will imrcxluce the electron microscope and
thechangesthishasbroughttothesrudyofcellstructure.
Light microscopy
We use microscopes ro magnify the cells of biological specimens in cnler ro see them at aU. Figure 1.4
showstwotype:sofligh!microscope.
In the simple microscope (the hand lens), a single biconv= lens is held in a supporting frame
so that the instrumem can be held do.se ro the eye. Tcxlay a hand Jens is used ro observe =temal
structure. However, some of the earliest detailed observations of living cells were made with
single-lensinstruments(seeFigurel.2)
using th e
compound
m·croscope
turre1-as ;1tstumedtheobject"""
dkkintoplace.fim!hemeDum-
power.!hen thehign-"°wer
condenser - foc:useslightonto
!heobjectwith iris diaphragm-
usedtova,ythefr,tensityollignt
,eachingtheobjea
Jn the compound micro scope. light rays are focused by the condense r on to a specimen on a
microscopeslideonthestageofthemicroscope. Lighttransminedthroughthespecimenisthen
focused by two sets of lenses (hence the name "compound" microscope). The objt.-ctin- lens
forms an image (in the microscope cube) which is then funher magnified by the t.')'C plece lens,
producingagreadyenlargedimage.
Cells and tissues examined with a compound microscope must be sufficiently transparent for
light rays ro pass through. When bulky tissues and pans of organs are ro be examined. thin sections
are rut. Thinsectionsarelargelycolourless.
1.1The microscope in cell studies I
Examining the structure of living cells
Livingrellsarena1onlytinybutalsotran.sparent. lnlightmicroscopyitiscornmonpracriretoadd
dyes or stains ro introduce sufficient contrast and so differentiate structure. Dyes and stains thm are
rnkenupbylivingcellsareespeciallyuseful.
p<pette Mottingpeper
sra;nd,awnacros,,
unde,thec,,.,e,s lip
Recording observations
What you see with a compound microscope may be recorded by d rowings of various types. For a
clear,simpledrawing:
• useasharpHBpencU and a de-an eraser
• use unlined paper and a separate sheet for each specimen you record
e drawcle-ar,sharpoudines:mdavoidingshadingorcolouring
• use most of the available space to show all the features observed in the specimen
e labeleachsheetordrawingwiththespecies.conditionsOivingorstained).transversesection
(fS)orlongitudinalsection(LS),andsoforth
• label your drawing fully, with labels positioned clear of the structures shown , remembering that
view (ph ase contrast) of th e layer of the The lini»g of the m,mac:h oomist, of columnar
ce lls (e pith e lium)liningth estomachwa ll epitheMum.Allcel ,secre!emucu,copiously.
a>"mM31
l'p<theliumcell
="'
"'""'= 0 0
"~'"' 0
0
-·-
bas<>mem
Ahcrna tivdy, images of cells and tissues viewed may be further magnified, displayed or
projected (and saved for printing out) by the technique of dig ital microscop y. A digital
microscope is used, or alternatively an appropriate video camera is connected by a microscope
coupler or eyepiece adap1or that replaces the standard microscope eyepiece. Images are
displayed via video recorder, TV monitor or computer.
Figure 1.7Measuringthesize
of cells
withi, i,
diaJ)h,agm
~--=-
bya lignme nt w iththesca leonthe
eyepi«eg raticule
thestagemkrnmete<isplac:edoothe
stageinflact'ol th.epreparedsjidearnl
exam,nedatthesamemagnilicatioo
=~,:~:~sr~.,..~ thell"ll',)SIJerrentoithered
bloodcel di,.,,.,.te,i,coo,,..,ted
1.5(1500~,)
' inthiscase.thelt'dblood
cellawea"toh..,ea
~
0 lOfl-lTI
d,.,,.,.1 .. ofaboo!81'ffi
Figure1 .8Record,r,gsizebymeamofsc:alebm
We use the ratio of these values to work out the 3crual length of the Amoeba.
100µ actu3llengthofthecell
19000µm • 95000µm
• SOO""'
(Note th3l Amoeba moves about by streaming movemerus of its cytoplasm. In this image the cell
is =tended and its length seems large, perhaps. It is equally likdy to be photographed in a more
spherical sh3pe, of diameter one tenth of its length here.)
9
magnificatiOfl • !~~~m • xl90
10
1.1The microscope in cell studies I
3 Finally. given the magnlfica1ion of an image. we ca n ca kulaie its r eal s ize.
Look at the images d the human cheek cell in Figure 1.3 (page 5).
5 Usingthemagnif1cation Measure the observed length of the cell in mm. It is90mm
given for the Conven this length to µm· 90 mm • 90 x 1000 µm - 90 OOOµm
photomicrograph of the
celloff/odeainFigure Usetheequauon aauals,ze(A) - m:;::::;:n(~M) - 90000 - 1125µm
1.3{pageS},calculate
the actual length of This size is greater than many human cheek cells. It suggests these epithelial cdls are squashed
the cell flat in thepo:sitiontheyhaveintheskin, perhaps.
.,
a irloc k/spec imen port
tfiespecimenisintroducedwithout
tfiel=ofYMUUm
obje ctive
electmmagneticlensthatfocuses
thefif,t image(Mcordir,gtovoltage)
proj ector
electrnmagneticlensth~tmagnifies~
part oftfiefi"t image
viewing port
withbinocula,Yiewe,
fluorescent screen
coatedwithelectron-sensitMecompound
Electrons are ne gatively cha rged a nd are camera cha mber
easily focused using e lec troma gnets. alk,ws~blackandwhrtephotog,aphK
imagetobemade(withthepossibility
transm·sson e lectron m· roscope of further magnifica tion)
Figure 1.10 Usingthetrammissiooelectroomicrosc"!"'
l2
1.1The microscope in cell studies I
2 Air inside 1he mlcro8CO!>C would d e flec t 1he deetrons :ind des tro y the beanL
lhe interior d the microscope must be under a varuum. Because of the varuum, fX> lid,,g specimens
8 Given the magnification can su,vive inside the dcctron microscope when in use. 'W-lter in cdls would boil away in a vacuum.
oftheTEMofalivercell As a result, before observations are possible, a specimen must have all the water removal.
inFigure1.11,@lrulate Sections are completely dehydrated . This has to be done whilst keeping the specimen as 'life-like'
a thelengthofthecell in structure a., ispos.sible. Thisisachallenge, g iventhatcellsareoo-9()percentwater. his after
b thediameterofthe the removal d water that the sections have the electron-dense stains adda:I.
nucleus
lhe images produced when this type of section is observed by the electron microscope are
calledtran.smissionelectronmicrographs('ITM)(Figurel.11).
13
D cellstructure
rep licaoffreeze~tchedsurface
Figu re 1.13 Trans mission electron mic:rographs from th in-secbor.ed ar>d freeze-etched mater;al
14
1.2 Cells as the basic uni~of living organisms I
Cytosol. organelles and the cell surface membrane make up a cell - a unit o f structure and function
which is remarkably able to survive, prosper and replicate itself. lbe molecules present in cells and
how the chemical reactions of life are regulated are the subject of Topic 2. lbe structure of the cell
membrane and how molecules enter and leave cells is the subject of Topic 3.
The structure and function of the o rgandies is what we consider next. Our understanding
of organdies has been built up by examining TEM.s of very many different cells. The outcome, a
detailed picture of the ultrascructure of animal and plant cells, is represented di3gmmmatically in a
generaliseclcellinFigurel.14.
plant cel l
tempor'")'vao.oles
formedbylnlood"]
~JU'fflamembrane
Figure1.14Theultra,t,uctu,eoftf>eeuh<y<>toc~nimalandplantcell
IS
D cellstructure
3 Endoplasmic reticulum
The endoplasmic reticulum consists of a necwork of folded membranes formed into sheets, tubes or
sacs that are extensively interconnected. Endoplasmic reticulum 'buds dr from the outer membrane
of the nuclear envelope, to which it may remain attached. The cytoplasm of mernbolically active
cells is commonly packed with endoplasmic reticulum. In Figure 1.15 we can see there are two
distinct rype., of endoplasmic reckulum.
• Ro u g h e ndo plasm ic n,tkulum (RER) has ribosomes 3ttached to the outer surface. At its
margin. \'Csiclcs are fonned from swelHngs. A vesicle is 3 small. spherical organelle bounded by
a single membrane, which beccmes pinched off 3S they separate. These tiny sacs 3re then used
to store and transport substances 3round the cell. For example, RER is the site of synthesis of
proteins that are 'packaged' in the vesicles. These vesicle, then fuse with the Golgi 3ppararus 3nd
3rethenrypicallydiscl,3rgedfromthecell. Digestiveenzymcs3redisch3rgedinthisw3y.
• Smoo th c ndo pla.~m ic reticulum (SER) has no ribosomes. SER is the site of synthcsis d
subsrnnces needed by cells. For example, SER is important in the manufacture d lipids and ste-
roids, and the reproductive hormones oestrogen and testosterone. In the cytoplasm of voluntary
muscle fibres, a speci3l form of SER is the sire d storage d calcium ions, which have 3n impor-
tant role in the contraction of muscle fibres
16
1.2 Cells as the basic uni~of living organisms I
SER a nd RER in cytoplasm, s howing o rigin from ou ter me mbra ne of nucl e us
o~:e:::~=~
QQ f0<exportfromcell
0
Cl
,rteofst0<ageof calciumions
smoothendoplasmk in(relaxed)volunta,ymusde
reticulum
4 Golgi apparatus
The G olgi appan,tus (Golgi body, Golgi compl=) consists of a Slack -like collection of flattened
membranous sac,. One side of the stack of membranes is formed by the fusion of membranes of
vesicles frcm RER or SER. At the opposite side of the stack, vesicles are formed from swdlings at
themarginsthatagainbecomepinchedoff.
The Golgi aPP"rarns occurs in all cells, but it is especially prominent in metabolically active
cells - for example, secretory cells. More than one may be present in a cell. lt is the site of
synthesis of specific biochemicals. such as hormones. enzymes or others. Here specific proteins
may be activated by addition of sugars (forming glycoprotein) or by the removal of the amino
add, methionine. These are then packaged into vesicles. ln an imal cdls these vesicles may form
lys.-.m cs . Those in plant cells may contain polysaccharides for cdl wall formation.
17
D cellstructure
Figure 1.16ThestmctureoltheGolgiapparatus
5 Mitochondria
Milochondrla appear mostly as rod-shaped or cylindrical organelles in electron
micrographs. Occasiconally their shape is more variable. 1bey are relatively large organelles,
typically 0 .5---1.5 pm wide, and 3.0-10.0 pm long. Mitochondria are found in all cells and
are usually presem in very large numbers. Metabolically very aaive cells will o:intain
thousands of them in their cytoplasm - for example, in muscle fibres and hormone-
secreting cells.
lbe mitochondrion also has a double membrane. The outer membrane is a smooih
boundary, the inner is infolded to form cris t:u,. The interior of the mitochondrion is called
the malrix . It o:intains an aqueous solution of metabolites and enzymes, and small circular
lengths of DNA, also. The mitochondrion is the site of the aerobic stages of respiration and
lnthemitod>ood<loomaovoftheeozymesof
feS!llra!loo~rehoused. and the"ene,gyc..-rer,cy" the site of the synthesis of much ATP (SC<."' below)
m,..._,desadeoo<ioelflphospl,all'{A.TJ?a,eformed Mitochondria (and chloroplasts) contain ribosomes , too. They appear as tiny dark dots
in the matrix of the mitochondria, and are slightly smaller than the ribosomes found in
the cytosol and anached to RER. The sizes of tiny objects like ribosomes are recorded in
Sved:>ergunilS(S). Thisisameasureoftheirrateofsedimentationincentrifugatior,,rather
than their aaual size. The smaller ribosomes found in mitochondria are 70S. those in the
rest of the cell OOS. We rerurn to the significance of this discovery later in the topic (the
endosymbiotic theory, page 25)
6 Ribosomes
Ribosomes are minute structures, approximately 25nm in diameter. They are built of two
sub-units, and do not have membranes as pan of their structures. O,entically, 1hey consist
of protein and a nucleic acid known as RNA (ribonucleic acid). lbe ribosomes found free
in the cytosol or bound to endoplasmic reticulum are the larger ones - classified as OOS. We
have already noted that those of mitochondria and chloroplasts are slightly smaller - 70S.
Ribosomes are the sites where proteins are made in cells. lbe structure of a ribosome
is shown in Figure 6.13. page 121, where their role in protein synthesis is illustrated.
Many different types of cell contain vast numbers of ribosomes. Some of the cell proteins
produced in the ribosomes have structural roles. Collagen is an example (page 48). A
great many other cell proteins are enzymes. These are biological catalysts. They muse the
Figure 1.17Thestructureofthe
mrtochondrion reactions of metabolism to occur quickly under the conditions found within the cytoplasm
18
1.2 Cells as the basic uni~of living organisms I
7 Lysosomes
Lysosomes are tiny spherical vesicles bound by a single membrane. lbey contain a concentrated
10 Explain why the mixture of 'digestive" enzymes. These are correctly known as hydrolrtic enzymes. They are
nucleus in a human producedintheGolgiappararusorbytheRER.
cheekcell(seeFigure
Lysosomes are involved in the breakdown of the contents of "food' vacuoles. For example,
1.3, pageS}may
harmful bacteria that invade the body are taken up into tiny vacuoles (they are engulfed) by special
beviewedbylight
m1croscopyman white cells called macrophages. Macrophages are pan of the body's defence system (see Topic 11).
appropriately stained Any foreign matter or food particles taken up into these vacuoles are then broken down . This
cell but the ribosome; occurs when lysosomes fuse with the vacuole. lbe products of digestion then escape into the
liquid of the cytoplasm. Lysoscmes will also destroy damaged organelles in this way.
When an organism dies, the hydrolytic enzymes in the lysosomcs of the cells escape into the
cytoplasm and muse self-digestion, known as autolysis.
Figure1.18Thestructweandfunctionofl,,sosomes
9 Chloroplasts
,~-
O,loroplasts are large organelles, typically biconvex in shape, about 4--IOµm long and 2- 3 µm
wide. They occur in green plants, where most occur in the mesophyll cells of leaves. A mesophyll
Figun,1.19Thestructu,eolthe
cellmaybepackedwithSOormorechloroplasts. Phorosynthesisistheprocessthatoccursin
chloroplastsandisthesubjectofTopic 13.
19
D cellstructure
Look a! the chloroplasts in the TEM in Figure 1.20. Each chlco-oplast has a double membrane. The
outer layer of the membrane is a continuous boundary, but the inner layer becomes in-tucked
to form a system of branching membranes called lamellae or 1hy lakokls. Jn some regions of the
chloroplastthethylakoidsarearrangedinflatteneddrcularpilescalled gran:,(singular: g r.rnum).
These look a Hnle like a scack of coins. It is here that the c hlo r o phyils and other pigments
are located. There are a large number of grana presem. Between them the branching thylakoid
membranes are very loosely aminged. The fluid outside the thylakoid is the s trom., , which
contains the chloroplast DNA and ribosomes (70S), together with many enzymes.
O,loroplasts are the site of phowsynthesis by which light is used as the energy source in
caibohydrate and ATP synthesis (see below)
O,loroplasts are one of a larger group of organelles called p las tld s. Plastids are found in
many plant cells but never in animals. lbe other members of the plastid family are leucoplasts
(colourless plastids) in which starch is stored. and chromoplasts (coloured plastids), containing
non-photosynthetic pigments such as carocene. and occurring in flower petals and the rooc tissue
of carrots
20
1.2 Cells as the basic uni~of living organisms I
11 WehaveseenthatanalysesofTEMimagesofcellshaveplayedanimportantpartinthe
disc:overy of cell ultrastructure. Carefully examine the image of a green plant cell 5hown
intheTEMinFigurel.21.UsingtheinterpretivedrawinginFigurel.11 as a model and
followingtheguidelinesonbiologicalrecordingonpage8,drawandlabelarepresentatKlfl
ofthegreenplantcellshowninFigurel.21
ATP, its production in mitochondria and chloroplasts, and its role in cells
ATP (aJcnoslne trlphosphate) is the universal energy currency molecule of celb. ATP is
formed from adcnoslnc dlphosphatc (ADP) and a phosphate ion (Pi) by the uansfer of energy
from olher reaaions. ATP is referred to as 'energy currency' because. like money, it can be
usedindifferentcontext'l.anditisconstandyrecyded. ltoccursincellsataconcentrationof
O.S---2.5 mg cm-J. ATP is a relatively small, water-soluble molecule, able to move easily around cells,
that effectively transfers energy in relatively small amounts, sufficient to drive individual reactions.
pho<phate
adenosfrieuiphosphateATP
adeooo;inediphospha te ADP
adenolinemonopho<phateAMP
m«>bcUcfh<"Om1 1<>dmeabollcp,oc••• -
'.'1,,:.:;::.::;;"'..,.:"'.:·:!~
•bull<11oemc1.:c...,.1,,,1<.,1,.,""""""c:1_..
21
D cellstructure
All' is a nucleotide with an unusual feature. It carries three phosphate groups linked together in a
linear sequence. All' may lose both of the outer phosphme groups, but usually only one at a time
is lost. ATP is a relatively small, soluble co-ganic molecule.
ATP conrnins a good deal of chemical energy locked up in its structure. What makes All' special
as a reservoir d stored chemical energy is its role as a common intermediate between energy-
}ielding reacrions and energy-requiring reacrion and processes .
• Energy-yielding reacrions include the phomphosphorylation reacrions of photosynthesis, and the
reacrions of cell respiration in which sugars are broken down and oxidiSN.
• Energy-requiring reacrions include the synthesis of cellulose from glucose, the synthesis d
proteins from amino acids, the contracrions d muscle fibres, and the acrive transport of certain
molecules across cell membranes. fco- example.
• lbe free energy available in the conversion of All' to ADP is approximately 30-34 kJ moi-' ,
made available in the presence of a specific enzyme. Some of this energy is lost as heat in a
reacrion, but much free energy is made available 10 do useful ,vork, mco-e than sufficient 10 drive
atypicalenergy-requiringreacrionofmetabolism.
• Sometimes ATP reacts with water (a hydrolysis reacrion) and is converted ro ADP and Pi. Direcr
hydrolysis d the terminal phosphate groups like this happens in muscle contracrion, for example.
• l>fostly, All' reacts with other metabolites and forms phosphorylated intermediates. making them
more reacrive in the process. lbe phosphate groups are released later. so both ADP and Pi
12 Outline why ATP is
an efficient energy
In summary. ATP is a molecule universal 10 all living things: it is the source of energy for chemical
currenc.ymolecule
changeincells.tissuesandco-ganisms.
22
1.2 Cells as the basic uni~of living organisms I
Insununary,thekeysirucruralfearuresoftypicalprokaryoticcdlsasseeninatypical
•srrucl\Jfe'.lthatocrnr
in a l bacteria
~ banie<aam~ which
al nutrientsillldwaste
pmdua,mu,1pass
ribosom... •- siteof
pmteinsymr.esk
nud80 id •- gene!k
:'!~~
ofaboul4000gelll'5
cell wa ll •- protl'Cts
cell fromruptu,e
causedbyosmosi,<>00
po:ss;bleharmlmm
otr>er0<ganisms
e leotronm ic rograph of£sdierichiaco/i
Figure1.23Thestructu<eof&heridii<>roli
Prokaryotes, e.g. bacteria, cyanobacteria Eukaryotes. e.g. mammals, green plants, fungi
cells are extremely small. typically about 1- 5 µm in d iamete, cells a re larger. typica lly 50--1 50µm
ntKleus absent d,ru!a, ONA helix in the cytoplasm. ONA ntKleus has d~tinct n tKlea. envelope {with p ores). with eh,~- of li r,ear ONA
notsupportedbyhistooeprotein helixsupportedbyhisto<>eprotein
cell wall p resent (made of pepbdoglycoo - kmg moleru!...s eel wall present in plants (largely of CPlulose) and fungi (laigely of the P™l'=hafide chitin)
ofam;no.Kidsandsugars)
few o«Jaoell,,s; membran oos structures ahsent many o,ga<>elles bounded by double membrar,e {e .g. chloroplasts. mitochondria. ntKleus)
Ofsingle membrar,e (e.g. Golgi apparatus. fysosomes. vi>CUOles. endoplasmic: retkulu m)
proteirissynthesisedinsmalltibosomes(70S) p roteirissynthesisedinlargeribosomee;{BOS)
cells have simple flagella cellshaveciliaorflagella.200nmindiamete,
some can fix atmospheric: nitrogen gas for lJSI' in the none can metaOO lise atmospheric: nitrogen gas but instead req uire nitrogen a lready
production of amino acids fo,pmtein synt hesis combined in moleru!...s in orde, to make proteins from am ino .Kids (pa.ge 120)
23
D cellstructure
pili
cellsurlace
memb<ane capsule(polysacchilfKle
coat)asitdditiooal
cytoplasm prntectioo
Figure1.24ThestructureofabactNium
24
1.2 Cells as the basic uni~of living organisms I
A possible origin for mitochondria and chloroplasts
Pre.sent-day prokaryotes are similar to many fossil prokaryotes. Some of these are 3S<Xl million
years old. By comparison, the earliest eukaryote cells date 1:xick only 1000 million years. Thus
eukary(){esmusihave evolvedsurroundedbyprokary<Xes,manythatwerelong-esrnblished
organisms. How the eukaryotic cell arose is nOl known. However, there is evidence that
the origin of the mitochondria and chloroplasts of eukaryotic cdls was as previously
independent-living prokary{){es.
It is highly possible that. in the evolution of the eukaryotic cdl, prokaryotic cdls (which at
one stage were taken up into food vacuoles for digestion) came to survive as organe lles inside
the host cell . rather than becoming food items! If so. they have become integrated into the
biochemistry of their 'host' cell, with time.
Theevidenceforthisoriginisthat mitochondriaandchloroplastscontain:
• a ring of DNA, like the circular chrcmosome a bacterial cell conrnins
• small (70S) riboso nu,s, like those of prokaryoies
The present-day DNA and riboscmes of these organelles siill function with roles in the
synthesis of specific proteins, but the mitochondria and chloroplasts themselves are no longer
capable of living independently.
It is these features that have led evolutionary biologists to propose this cndo,.-ymblolk
1lu,Dr)· of the origin of these organelles ('cndo' • in.side, 'symbiont' • an organism living with
another for mutual benefit)
RNAviru""
1 5'ngle-manded 2doobie-manded
~,,:host, re<Mrusol
""'"""M<ts
@so,~:25nm
""-
immuoodefkiency
Wus,1-HV(retr<Mru,)
Figure1.25Adas,ificationofviruses
25
D cellstructure
~~
pro1eincoa1(capsid)ol
~ :!r;::;·
~tidebuidongbkrls
5' ::::~:1:~=~~o~,~~~
in farted
laaf
Figure1.26Toba<:eomosakvirus
26
1.2 Cells as the basic uni~of living organisms I
........., -
_,
~!...1
organi"'tioo
olliving!hingl
aoss-Sl'Clioool
>---- giant"'""' cl'II
otsqu,d
Histology
studyofti,,..,...
bychemi::alanalysi,
Figure1.27Sjzerelationshipsonalog.lrithmicsc:ale
27
D cellstructure
• Cellsarethe buildingblocksoflivingthings. They are derived • Examination of tran smission e lectron microg raphs has
fromothercellsby divis ion andtheyarethesiteofallthe revealedthatthecytoplasmoftheeukaryoticcellcontains
chemical reactions of life (metabolism}. A (ell is the smallest numerous organelles, wrne about the size of bacteria,
unitoforganisationwe(ansayisalive suspendedinanaquaticsolutionofmetabolites{calledthe
• Cells are extrem e ly sma ll. They are measured in units of a cytosol)surroundedbythecellsurfacemembrane
thousandths of a millimetre (a micron - µm) and they must be • Manyoftheorganellesare membran e -boundstru ctures,
viewed by microscopy. In the laboratory we view them by light including the nucleus, mitochondria, chloroplasts, endoplasmic
microscopy using a compound microscope reticulum,Golgiapparatusandlysosomes.Theorganelleshave
• The(ellsofplantsandanimalshavecommonfea tures, specilicrolesinmetabolism.The biochemicalro lesofthe
including a nudeus, cytoplasm and cell membrane. To observe o rganell esareinvestigatedbydisruptingcellsandisolating
andresolvethedetailedstructureswithinthecytoplasm, theorganelleslorlurtherinvestigation
e lectron microscopy is required. The distinctive features • Virusesarenon-cellularstructuresthatcomistofacoreof
ofplantcellsareacellulosecellwall,thepresenceoflarge nucleicacid {DNA or RNA}surroundedbyaproteincoat,
permanentvacuolesandthepossiblepresenc:eofchloroplasts, called a caps id. They are extremely small when compared
the site of photosynthesis withbacteria,andtheycanreproduceonlyimide1pecif1c
• Thesimplestcellularorganisationisshownbybacteria.Here livingcells,sovirusesfunctionase ndopara sitesintheirhost
thereisnotruenucleus.Theseunicellularorganismsare
c.alled prokaryotes.Thecellsofplants,animalsandlungi
arelargerandtheyhaveatruenuc:leus.Theselivingthings
arecalledeukaryotes
28
AS Level
2 Biological molecules
Thistopicintroducescarbohydrates, proteinsandlipids:organic Th eemphasisinthistopicisonth erelationshipbetwee nmol ec ula r
moleculesthatareimportantince lls.Nucleic.: ids arecoveredin a structuresandtheirfunctions.Someoftheseideasarecontinued
separate topic. Biological molecules are based on the versatile element in other topics, for example, th e functions of haemoglobin in gas
carbon. This topic explains how mac romolecules. w hich have a great tra ns port in Transport in mamma ls, phospholipid s in membranes in
diversity of function in organisms. a re assembled from smaller organic Cell me mbra nes and tra nsport a nd antibodies in Immunity.
moleculessuch asg lucose,a mino acids.glycerol a ndfattyacids.
Life as we know it would not be possibl e without wat e r.
Und erstandingthepropertiesofthisextraordin ary molecule is
a n esse ntia lpa rtof a nystudyofbiologica lmolecules .
Teo;tsforbiologic.al
moleculesrnnbeu1edina
By the end of this section you should be able to:
varietyofcontexts,suchas a) carryouttfflsforreducingsugarsandnon-re ducingsugars,th e iodine inpotassiumiodide
identifying the contents of solutiontestfor,tarch,theemulsionte<tforlipidsandthe biure ttestforproteinstoide ntifythe
mixtures of molecules and
following the activity of b) carryout a semi-quantitative Be ne dict'stestonareducingsugarusingdilution,standardising
digestive enzymes
th e testandusingthe results(colourstandardsortime tolirstcolourchange)to e<timatethe
29
El Biological molecules
Carbohydrates and lipids By the end of this section you should be able to:
have important roles
in the provision and a) describethe ringformsofa-glucose andp..glucose
storage of energy and b) define the terms monome r, polym er, macromol ec ule, monosaccharide, dinccharide and
foravariety ofother polysaccharide
functiomMJc:hasproviding c) describethe formationofaglycosidkbondbyconde n,ation,withrefe rence bothto
barriersaroundcells:the
polysaccharid.,., andtodi,accharid.,.,,indudingsucrose
phospholipidbilayerofall
d) describethe breakage ofglycosidkbondsinpolysaccharidesanddi,accharid.,., byhydrolysi,,with
cell membranes and the
refe rence tothe non-reducing,ugartest
cellulosecellwallsofplant
cells e) describe the molecularstructureofpolysaccharide,includingstarch(amylcse andamylopectin),
gly,:oge nandcellulose andr elate th.,.,e rtructur.,., toth eirfunctionsinlivingorganisms
f) describethe molecularstructure ofatrigly,:e ride withrefe rence tothe formationof e,te rbonds
andrelate the structur e oftrigly,:erid.,. totheirfuncticn,inlivingorgani,m,
g)describethe structure ofaphospholipidandrelate the rtructure ofphospholipid,totheir
functions in living organisms
30
2.2Carbohydratesandlipids l
Carbohydrates
Carbohydrates are the largest group of organic compounds. 1bey include sugars, srnrch, glycogen
and cellulose. Carbohydrates are substances that contain only three elements carbon, hydrogen and
oxygen. lbe hydrogen and oxygen atoms are present in the ratio 2:1 (as they are in water, H,O). In
fact, we represent carbohydrates by the general form ula c.£11.P)y.
We stan by looking at the simplest carbohydrates
~
thetwofo,msolglucose
andtakesaringorcydicform. This is its
s trucmral form ula. lbe ring doses up
depend onthepositionsof whentheoxygenoncarbon-5anachesitself
the - H and -OH attached rocarbon-1. Theglucosering, containing
~11 lucose
tourboo-1whenther ingdoses fivecarbonatomsandanoxygenatom,is
called a p yra n osc rlng. Funherrnore.the
~or simplicity and conven>l'Jlce P}TaJX>Seringexists in two forms. These are
itistheske~l/om,ulaethat the a-form and the ~form. depending on
aremostfrequentlyusedin whether a - H atom was trapped 'up" (a-form)
recording biochemical reactions or 'down" (~-form) when the ring closed. So,
and,howingthestructureof
bk,logical!y..ctivemolecules there are two forms d glucose, known as
oln-<J lucose of ~-<J lucose a -glucooeand ~-glucooe(Figure2.2).
Figure2.2Thestructureofn-<Jlurnseand~-glucose
"
El Biological molecules
Incidentally. fruc toiscisanOffierhexosesugarfoundincells. hhasaringsiructuredifferentfrom
that of glucose. Fructose form., into a ring with four carbon atoms and an oxygen atom (known as
fur:mose ring). Fructose is by far the sweeteSI common sugar: fructose is used in the food industry
inthemanufacrureofsweetsandvariousconfectioneries
Som e function a l
g roups o f s ugars
akoholgroop
"I
- C- 0-H
H ~1-+,0H O H hydrnxylgroup - OH
aldel,yde H
)c(~Hc- c~)c( ~~:~t'"'-<'o
OH OH
p suffy,J,ylgroup - S- H
carboxyl group - (OOH or - C'o- H
a-glucose
analdme
l 6H (or - CHO)
\
carbonylgrnup C=O (thisgrnupispartof
I theaklel,yde.keto
andc arboxylgroop,)
32
2.2Carbohydratesandlipids l
3 Finally, place 10cm1 of dislilled water in a test tube (Tube I) and IOcml d a l(m sucrose
solutioninafinalte.sttube(fube5).
4 Add 5cm1 of Benedict's solution to each tube.
5 Placeallfiverubes inthewaterbath for5minutes,gendyagitatingthemduringtheprocess.
Finally, remove the tubes to a rack where any colour changes can be observed (Figure 2.4)
5 cm' of Beoedict's solution - test tubes were placed in a boili ng ---- tubes were tram/erred to a ,ad and the
{blue)was added to 10 cm' wate, bath fm 5 mfr, utes colours compared
ofso lubootobetested
with sucrose
solution
Figure2.4Thetestforieduongs.uga,
33
El Biological molecules
making aserialdilutio nofglu cosesolution
~ 6 discard2cm l
_ D D D D D lr-----;:Jofthesoltrtionintube6
1 2 3 4 5 6
2add2cm1 wate,totubes2---6
Length of Molecular
carbon chain formula
a) fl,C.,:::P
respirabonand
H- ~ - 00 pl,otmynthesis
CH 10H
deoxyribose
2 1..istthepropertiesofthe
carbon atom responsible
forthehugearrayof
organic compounds.
2.2Carbohydratesandlipids l
Disaccharides
Disaccharides are carbdlydrates made of two monosaccharides combined together. An example is
Figure2.6Suc:mse{a (I-glucose
dis.icchafide)andthe
mOI\OS.Jccharidesthatlom,it
+ Hp hy<lrolysi1
water condensation
35
El Biological molecules
lbe-re are ocher disaccharides formed in cells, depending Oil which -=ccharides are involvn:l
and whether they are in their a or 13 condition. Two other commonly occurring disaccharides are
m3lto.seandlactose.
Jl.fallt>lie (a -glucose + a -glucose) is a product of st:irch hydrolysis. The =traction of m3Jto.se
from germinating barley (a., m3Jt extract) is an important industry. Mah is used in brewing and in
OH 1--10
Polysaccharides
Polysaccharides are built from very many mOflO<Saccharide molecules condensed together.
Oncetheyarepanofalargermolecule,theyarecalledresidues. Eachresidueislinkedby
glycosidic bonds. 'Pol)! me:m.s "many' and, in fact, thous:mds of 'saccban"tte re.sidues make up
a polysaccharide. So a polysaccharide is an example of a giant molecule, a m 3cromolecu le.
Normally each polys:,ccharide contains only Ofle type of mon o m er. Cellulose is a good =ample -
built from the monomer glucose
Bonds in polysaccharides
Atoms narurally combine together (they "bond") to form molecules in ways that have a stable
a1Tangement of electrons in the outer shells of each atom (see Appendix 1 Oil the CD). Jn
covalent bon di n g, electrOfls are shared between atoms. Covalent bonds are the strongest
bond.s in biological molecules. In polys:,ccharides, the individual monomers are held by covalent
bonds.
However. entirely different bonds, called h ydroge n b o nd .. , are also fonned. We can illustrate
how hydrogen bonds form by reference to the water molecule.
Water is composed of one atom of oxygen and two atoms of hydrogen, also combined by
covalent bonding. The water molecule is triangular rather than linear. and the nucleus of the
oxygen atom draws electrons (negatively charged) away from the hydrogen nuclei (positively
charged) - with 3n interesting consequence. Althou gh overall the water molecule is electrically
neutral,thereisanetnegativechargeOfltheoxygenatom(conventionallyrepresentedbyGreek
letter delta as 1)) and a net positive charge Oil the hydrogen atoms (represented by .S). Jn other
word.s , the water molecule ca1Ties 3n unequal distribution of electrical charge within it. This
unequaldistributionofchargeiscalledadipo le . Moleculesthatcontaingroupswithdipolesare
known as polar m o lec ules . Water is a polar molecule (see Figure 2.27, page 56).
36
2.2Carbohydratesandlipids l
Dipoles exist in many different molecules, especially where there are -OH, ----c=o. or =N-H
groups, so hydrogen bonds can fonn between these groups. Hydrogen bonds are weaker
than covalent bonds. but never the Jess are very importam in the Slructure and properties of
caroohydrates (including polysaccharides) and proteins. For example, molecules containing these
groups are attracted to water molecules because of their dipoles. and are therefore said to be 'water
loving'or hydrophilk.
Cellulose
Cellulose is by far the most abundant carbohydrate it makes up more than SO per cent of all
org:mic carbon. (Remember. the gas carbon dioxide. co,, and the mineral calcium carbonate,
CaC0 3, are examples of inorganic carbon.)
Cellulose is a polymer of 13-glucose molecules combined together by glycosidk bonds becween
carbon-4ofoneJ3-glucosemoleculeandcarbon- 1 ofthenext.Successiveglucoseunit.sarelinked
at 100° to each other (Figure 2.8). This structure is stabilised and strengthened by hydrogen bonds
between adjacem glucose units in the same strand and, in fibrils d cellulose, by hydrogen bonds
between parallel strands, too. In plant cell walls adclitional strength comes from the cdlulose fibres
beinglaiddowninlayersrunningindifferemdirections.
The chemical test for cellulose is that it gives a purple colour when Schultz" solution is adclcd .
The cell walls d green plants and the debris d plants in and on the soil are where cdlulose
occurs. ltisanextremclystrongmateri.al - insoluble,toughanddurablebutslighdyclastic.
Cellulose fibres are straight and uncoiled . When it is =tractn:I from plants, cellulose has many
industrial uses. We use cellulose fibres as conon, we manufacture them imo paper, rayon fibres for
clothcsmanufacture,nitrocelluloseforcxplosivcs,celluloseacctateforfibresofmultipleuses,and
cellophane for packaging.
p--glurnsemolecules
twop..glucosemolecuk-i rotated180° with
I \
:-*
andtheformationofarespecttotheothe~thestrandsareheklstraightbyc°"alentbondsbetween
1.4g:=osid~tk CH,OH glucoseun;ts.anldbyhydrogenbondswithjnthestrand
, :-V?o*'"'':o~o
OH~ - ~ ~- OOH
0
yJ.oJ,1
H
l~H
~-
/,_.!
OOH
;;i_j
l!Ai
~OH
OH~- f ~
~'":
....
o
00
O
v'"'': 00
o j" O
00
cellulose strand,
packed together lecules
to form fibrils ht.
nchedchains
cellulose laid
down at different
electron microg raph of angle,
cellulose in a plantcellwall
()1 1500)
Figurel.8Cellubse
37
El Biological molecules
Starch
Starch is 3 mixture of two polysaccharides (Figure 2.9).
• Amyl,-, is an unbranched chain of several thousand 1,4 linked a -glucose units
• Amylopcc tin hasshonerchairuofl,4linkeda-gluo:iseu n itsbut, inaddition,therearebranch
points of 1.6Hnks along its chains
Thesecov:ilentbondsbetweenglucoseresiduesinsmrchbringthemoleculestogetherasa
4 Starch is a powdery helix . The whole starch molecule is stabilised by countless hydrogen bonds between parts of the
material;cellulose component glucose molecules.
i1a1trong,f1brou1 Starch is the major storage carbohydrate of most plants. It is laid down as cornpaa grains in
sub51ance;yetbothare pla.stids called leucoplasts. Smrch is an important energy source in the diet of many 3nimals,
madeofglurose.What too. Its usefulness lies in the compacmess and insolubility o f its molecule. However, it is readily
features of the cellulose hydrolysed to form sugar when required. We sometimes see 'soluble starch' as an ingredient of
moleculeaccountforitl m:,nufacrured focxls. Here the smrch molecules have been broken down into short lengths. making
strength7 them more easily dis.solved.
··. ..,,
. /
"f , t..
~~ ~ !%<!arch
solutloo
0.1%ru«:h
solU!loo
0.01%stafeh
,olutkln
a) Tes!ooapotato lubefrutSllrlace b)Testoostarchsollltioosofararqeofconceotratklns
38
2.2Carbohydratesandlipids l
5 Definetheterm1
'monomer'and
'polymer', giving Lipids
examples from the Lipids contain the elements carbon, hydrogen and oxygen, as do carbd,ydrates. However, in lipids
carbohydrates. theproportionof=ygenismuchless.Lipidsareinsolubleinwater.Infacttheygenerallybehave
as 'water-hating" molecules, a propeny described as hyd rophobic. Lipids can be d issolved in
organicsolventssuchas alcohol(for=ample,ethanol),propanone andecher.Thispropenyis
made use of in the emulsion test for lipids (Figure 2.13)
Lipidsoccurinlivingthingsasanimal fatsandplantolls. Theyarealsopresentasthe
phos pholipids of cell membranes. In addition, there are Ol:her, more unusual forms of lipid. For
example, the steroids from which many growth and sex hormones are produced are lipids. The
waxesfoundonplantsandanimalsarealsolipids
39
El Biological molecules
The fatty acids present in fats and oils have long hydrocarbon ·rails'. Toe.... are typically of about
16 to 18 carbon atoms long but may be anything between 14 and 22 (Figure 2.12). The
hydrophobic propenies oftriglyceridesare duetothesehrdrocarbontails.
We describe fatty acid molecules as 'acids' because their fuoctional group (--COOH, carbCJxrl)
tends to ionise (slightly) to produce hydrogen ions, which is the propercy of an acid.
--.COOH s===' --.COO - + H~
ltisthecarboxr[functional groupofthreeorg:micacidsthatreactwiththethreehydroxr[
functional groups of glycerol to form a triglyceride (Figure 2.12). lbe bonds formed in this case are
known ester bonds .
"mo,,<om: (
:J
hy<Jrocatbootail
,7,7,7,7,7,7,7, !l ::=i::::
I
H,~/1'-~/1'-.~/1'-~/1'-~/)'-l/)'-l~,l/c\ o H HO - ~---H
!" !":.:.,.:,:."..:"""':"~~",":~:~
j '"d '"""'
abondisf0<medbet,,.....,,thecarhoxylgroup (- CCX)H)offattyac:id
andooeofthehydroxylgroups{---OH)ofgly<:erol,toproducea monog lyce ride
a~ ~ R
+ i -(CH,lnCH i - - - - - -- := t =: , : ~ : : ~ H i
i
glycerol fatty acid monoglyceride
"I 0II
H- C- 0 - C- (CHi),,CHi
I ~
H- C- 0 - C- (CHi),,CHi
I ~ The threefattyaddsina
H- 1 - 0 - C- (CH 1),,CHi =e:~;:.b;i~~!:t
40
2.2Carbohydratesandlipids l
lflsteam.meltlngpolmTJ."C
Figure2.14Satu,atedar>dunsaturatedfattyacid,.aOO!hetriglyceridestheylorm
Where several double bonds occur the resulting fat is called polyunsaturated. Fats with unsaturated
fatty adds melt at a lower temperature than those with saturated fatty acids, because their
unsaturated hydrocarbon tails do not pack so closely together. You can see why in Figure 2.14.
"
El Biological molecules
This difference between saturated and polyunsaturatn:I fats is important in the manufacture of
margarine and butter spreads, since the later perform better 'from the fridge'. There is disagreement
about the value (or a1her...ise) of 'polyunsaturates' in the human diet. What seems dear is that it is
bettertoeatlessratherthanmorefatinourdiets.
Electrical insulation
Myelin lipid in the membranes of Schwann cdls, forming the sheath around the axons of neurones
(page 313), electrically isolates the cell surface membrane and facilitates the o:induaion of the
nen,eimpulsethere.
Phospholipids
A phrupholipid has a very similar chemical struaure to a triglyceride, =cept that one of the fatty
adds is replaced by a phosphate group. The phosphate group is ionised and therefore water
soluble. Consequently, phospholipids o:imbine the hydrophobic propenies of the hydrocarbon tails
with the water-loving (h ydrophilic) properties of the phosphate group
A thin layer of phospholipid floats on water, forming a monolayer with the hydrophilic heads
in water and the hydrophobic tails projecting. With more phospholipid present, the phospholipid
molecules form a bilayer, with the hydrophobic tails aligned together. We return to this when
looking into the struaure of cell smface membranes (fopic -0.
42
2.2Carbohydratesandlipids l
6 Describetheproperties
giventoal ipidwhen
it(ombineswitha
phosphate group phospha te
Figure2.16Phospho1ipids
Other lipids
lfyoucomparethestructureof aste roid suchascholesterol(Figure2.17)withthat ofa
triglyceride(Figure2.14),youcanseejuschowstrucrurnllydifferentthelipidsare. The 'skeleton'of
a steroid is a set d complex rings of carbon atoms; the bulk of the molecule is hydrophobic but the
pobr hydroxyl group is hydrophilic. Steroids occur in boch pbnts and animals, and one widespread
form is the substance c holes tero l.
ro les:
O>olescerol is an essenti.tl component d membranes of pl.tms and an irn:,ls. In rn:,mn13ls, the sex
hormones proge.sterone, oestrogen and testo.sterone and also growth hormones are also produced
from it. Bile salts (involved in lipid transport front the inte.stine to the tissues, in the blood) are also
synthesised from cholesterol. Consequently. cholesterol is essential for normal, healthy metabolism.
However, =cess blood chole.sterol n13y cause harmful deposits in artery walls. causing a disease
called atherosclerosis
Finally. waxes are e.sters formed from a fatty add and a complex alcohol (in pi.tee of glycerol)
They are produced by plants and used as the waxy cuticle that prot"ects the plant stem and leaf
epidermis from loss of water by evaporation. The same role is fulfilled by the wax that insects have
on the outer surface of their cuticle. Bees use wax in the honeycomb cells they build for rearing
theiryoungOarvae)andstoringhoney.
43
El Biological molecules
AA understanding of protein By the end of this section you should be able to:
structureandhowitis
relatedtofunctioniscentral a) describethestructure ofanaminoaddandthe formationandb,eakageofapeptidebond
to many a ~ of biology, b) explainthe meaningofthe te rmsprimaryrtructure,secondarystructure,tertiarystructure and
!.Uc:hasenzymes,antibodies quaternarystructureofprote insanddescribethetypesofbonding(hydrogen,ionic,disulfld ea nd
and muscle contraction. hydrophobkinteractions)thatholdth.,..,mole culesin,hape
Globular and fibrous c) desc ribe the molecularstructu,eofhae moglobinasan exampleofaglobularprotein,andof
proteimplayimportantroles
collagenasan6ampleofaflbrousprote inandrelate these rtructurestothe irfunctions
in biological processes such
d) explainhowhydrogenbondingoccursbetween watermoleculesandrelate the p,opertiesof
asthetramportofgases
and providing support for wate rtoitsrolesinlivingorganisms
tisrues
Water is a special
molecule1\ithextraordinary
properties that make life
possibleonthisplanet150
million kilometres from the
Proteins
Proteins make up about two-thirds of the tocal dry mass of a cell. They differ from carbohydrates
and lipids in that they contain the el=ient nitrog e n , and usually the element sulfur. as well as
caJbon,hydrogenandoxygen.
Pro1eins are large molecules formed from many a mino add ,;; combined in a long chain
Typically several hundreds or even thousands of amino acid molecules are combined together
to make a pro1ein . (So a prolein can also be described as a m acrom o lec ule.) Once the chain is
conscructed, a prorein takes upa specificshape. lbeshapeofa protein is closely related to the
functions it performs. We will rerum to this aspect of protein structure, after we have =amined the
propertiesofindividualaminoacids.
Amino acids, as their name implies. contain two functional groups, a basic a m i n o grou p (- NHJ
and an acidic carboxyl gro up (---COOH). In the naturally-OCC"Urring amino acids both functional
groups are attached to the same carbon atom. Since the carboxyl group is acidic:
-COOH a===' -COO - + H+
andtheaminogroupisbasic:
- NH, + tt+ a===' - NH3
an amino acid is both an acid and a base. a condition known as a mpho icric. The remainder of the
molecule,thesidechainor R-group ,isveryvariable.
The bringing of amino acids together in different combinations produces proteins with very
different properties. This helps explain how the pro1eins in organisms are able to fulfil the very
different biological functions they have.
44
2.3Proteinsandwater .
7 Vv'hatdoesthe
symbol'R'represent
in a generalised
formula of an organic
compound?
~ - - - - - theR-<Jroupsof - - - - - H c CH
th ree i miooiKids l 'iff" l
8 Thepossible
numberofdifferent
l 'oH l 'oH
polypeptides(P}that
can be assembled is glyci ne
given by
Figun,2.18Thestructureofaminoac>ds
P = A"
whereA = the
number of different
The peptide bond
types of amino acids Two amino acids rnn react together with the loss of water to form a d ipeptide. The amino group
available and of one amino acid reacts with the carboxyl group d the other, forming a pe ptide b o nd (see
n = the number of Figure 2.19 on the next page). Long strings of amino adds linked by peptide bonds are called
amino acid monomers poly p e ptides. A pep!ide or pro1ein chain is assembled, one amino acid at a time, as we shall see
that make up the
polypeptide molecule
The terms 'polypeptide' and 'protein" can be used interchangeably but. when a polypeptide is
Given the naturally about 50 amino acid molecules long, ii is generally agreed to be have become a protein.
occurring pool of
20differenttypeJ;of
aminoac:ids,rnlculate Testing for proteins
how many different The blure t tes t is used as an indicator of the presence of protein because it gives a purple
polypeptides are
colour in the presence of peptide bonds (- C--N- ). To a protein solution, an equal quantity of
possible if constructed
sodium hydroxide solution is added and mixed. Then a few drops of 0.5 per cent copper([))
from S, 25 and SO
amino acid residues sulfate is introduced with gentle mixing. A distinctive purple colour develops without heating
respectively? (see Figure 2.20 on the next page).
9 Oistinguishbetween
condensation and The structure of proteins
hydrolysis reactions
The primary s tru c ture of a protein is the long chain of amino acids in its molecule. Proteins
Give an example of
,~h differ in the variety. number and order of these amino adds. In the living cell, the sequence of
amino adds in the polypepiide chain is controlled by the coded instructions stored in the DNA
of the chromosomes in the nucleus (Topic 6). Just changing one amino acid in the sequence of
a protein may alter its properties completely. This sort of ' mistake' or mutation does happen
(see page 125).
45
El Biological molecules
Figu,..,2.19Pept<lebood amino acids combine together. the amino gmup olooe with the e a ~ group al" the othef
formation
dopeptkle
f0<examl)le.~;neandalaninecan,eactlikethtl
• ,,o
tiutiftheamino,,-oupofrjydnereactswiththecarboxy!,,-oopol
alanine.aclitlerentpolypeptide.aOOyl.glycine.islormed
thn,eamjnoacids
comb<nelcxjether
tolormatnpepbde
0. 5%coppen:!Osu!fate
solution(...;adroppingbottle)
c;---;::::, D D lc::;-;i
purplecolou,
develops blue
46
2.3Proteinsandwater .
The secondary s trnctun, of a protein develops when pans o f the polypepikle chain rnke up
a panicular shape, immediately after formation at the ribosome (Figure 2.21). Paru of the chain
10 Describe three of the become folded or twisted, or both, in various ways. Two major structural forms are panirularly
type1ofbondsthat stable and common. Either pan or all of the pep!ide chain becomes coiled to produce an a -helix
maintain the tertiary
or it becomes folded into l!,.sh<.-.,1.s. These shapes are permanent. held in place by hydrogen bonds.
structureofprotefr11
The tc niary s truc tun, of a protein is the precise, compacr structure, unique ro that protein
that arises when the molecule is funher folded and hdd in a panicular compl= shape. This shape
is made permanent by four different types of bonding, established between adjacent pans of the
chain(Figure2.22)
lbe primary, secondary and tertiary structure of the pro-ein called lysozyme is shown in Figure 2.24
o:-helix {rod-like) ~sheet s
pmil,OOofam;ooaod
residuesandpeptidel;nka.ges
Figure2.21Thesec:oridary,tructureofproteins
hydrophobicintera ctions
T
flt="'
/
'<:Hi
andvande rW aalsforces
thesernme intoplay"tientwoorffiOfe
atom,areverydose (O.J---0.4 nmapart)
l
H"' l strongrnvafentbon.df=.J
bytheoxidationof - SHgroups
oftwocysletneside-chains
f=f,bood
weake!ectrostatkinteractioo
- CH 1- s- s- c between oppositely charged
,onscmayoftenbebrokenby
chang ,ngthepH
>---H-< R
>-m
m
1~ - o -1,--o--1i---c1-1i- rn,--NHi· --o- c-----<:H1-
electropositivej L e!ectronegative
hydrogen atom
Figure 2.22C,oss~inkingwithinapolypeptide
47
El Biological molecules
photomicrogra phofcollage<1 Fibrous and globular proteins
fibre{~OOO) - ma nytriple
he lices bound together Some proieins take up a tertiary structure which is 3 long strand, a nd are called fibrou s proldns.
They have structural roles in the body. Examples o f fibrous proieins are fibrin , a blood protein
involvedinthedoctingmechanism,and kcratin,foundinhair,hornandnails
Collagen
Collagen is the most abundant structural protein in animals in general. In our skin it forms a mat in
the deepes! byers. It occurs in tendons, cartibge, bone, teeth and in the walls of blood vessels. I!
also forms the cornea of the eye.
O,emically, a collagen moke ulc consists of three polypeptide chains, each in the sh3pe of a
helix. Each chain consists of about 1000 amino acids. 1bey are wound together as a triple helix
forming a stiff cable - a unique arrnngement in proteins. These helices are stretched out, meaning
this structure is not as tightly wound as an a -helix. This structure is Oflly able to form if every third
residue in e3ch polypepiide chain is glycine - the smallest amino add. In fact, 3long the helix a
rypicalrepeatingsequenceisglycine-proline-hydroxyproline.
The three helices are held together by numerous hydrogen bonds and have great mechanical
strength. Many of these triple helices He side-by-side, forming collagen fibres . Within the fibres,
-·-...
thec he mic a lbasisofth e
stre ngthof collage n collagen molecules are held together by covalent cross-linkages between the carboxyl group of
one amino add a nd the amino group of another. Also, the ends of individual collagen molecules
are staggered so there are no weak points in collagen fibres, giving the whole strucrure incredible
molea!lo>s,mlled
togethef!Olom, tensile strength. For example, aloadofatle-ast !Okgisneededtobreakacollagenfibrethatis
atripk'l>ek>: only I mm in diameter. This qualiry is essential in tendons, for example. Within the body, collagen
also binds strongly to ()(her substances, such as the calcium phosphate of bone. The strucrure of
collagenisillustratedinFigure2.23.
Other proieins take up a tertiary strucrure that is more spherical. They are called globular
proteins. Enzymes are typically globular proteins, including lysozyme an aruil:xicrerial enzyme
presentinanimalexcretionssuchastears(seeFigure2.24)
covalemtlonds
loon between
primary structure secondary structure
~-~
cl.Mns - togethef (thesequenceofamioo.tcids) (the,hapetakenupbypartsoftheamiooacodchain)
~maoyl¥Jrogen .,.,.....,,....,.,.,...""'~~
Figure2.23 Collagenan
exampleofafibrouspmte in ,.•..,1¥i-~·""
~~~"''"''-"':/,.
,_...,.._.,.-.,.•~
;""""#on."~"'1:
,~.... ,~ ....
t,_,.,,,,,,,cr,)
;;'""""-c,."",,,'..,
.. .. n._::::--
:::;,;:.:..1:
;:::.::::{
48
2.3Proteinsandwater .
Finally, the quaic rnary s!rucmrc of a protein arises when two or more proieins become held
togel:her, forming a complex, biologically aaive molecule. An example is haemoglobin. a globular
protein (globln) consisting of four pol~J>qXide chains (two a -chains and two IJ,-cha ins ). Each
polypeptide chain is associated with a non-protein hae m group, in which an Iron lon (FeH)
occurs (Figure 2.25). Since the pigment ' haem' is an imegral part of the qumemary protein and yet
is not itself made of amino acids, it is referred ro as a pros thetic group. A molecule of ox}·gcn
(Oz) combines reversib ly with each haem group in the 'pockei' where the iron ion occurs. This
reaction is dependent upon the panial pressun, of oxygen (high in the lungs. low in the
respiringtissuesofthebody).
Within the haemoglobin molecule the forces holding it together (and thereby maintaining the all
impon:am three-dimensional shape of haemoglobin) are inwardly-facing hydrophobic R-groups. On
the =terior of the molecule it is the presence of hydrophiHc R-groups that maintain the solubility
of the pro1.ein. Vast numbers of the resulting more co- less spherical haemoglobin molecules are
normally located in the red blood cells.
~~~
t
Eachs ubunit isacoojugated
protein.comistir,gofa
prnte;n chain (gkibin) attached
toaprostheticgmup (haem)
_,,;;:=- v\e;f, ::t::;::::::~~
, ~
1
,::'1,m
acompactlnrnecule
A prost heticg roupisa
"helper"molecule.enablir,g
othe,moleculestobe
biologka lly active
Haem isaflatmolecule of
fourpyrmlegroop,.held
~~: ·~n) ,,· '·,, c&"
togetherby=C- g10Up,: at
thecentreisanatom of;mn(II) ,'
Globin coosistsof150aminoiKffl,esidues in
thelom,ofahelixthatisfokledS--7times
Figure 2.25Haemogkibin:thequaternaryproteinofredcells
49
El Biological molecules
An imponam =cepcion to this arises in skklc-cdl a naemia . a genetically comrolled condition in
which Ofle of the amino acids that comprise the 13-chain is replaced (non-polar valine is substituted
for polar glutamic acid). The result is long fibrous haemoglobin molecules that distort and damage
the red blood cells and which hinders their efficient delivery of oxygen to the tissues. However. the
sickle cell condition may confer some resistance to malaria .
Denaturation of protein
Dcnamr:nion is the Joss d the three-dimensional structure of a protein. It happens when
the bonds that maintain the three-dimensiOflal shape of the protein molecules are changed
(Figure2.22).
We have noted that many of the propenies and uses of proteins within cells and organisms
depend on their particular shape. When the shape of a protein changes the pr(){ein may cease to
be useful . The biochemistry of cells and organisms is =tremely sensitive to conditions that alter
prote ins in this way.
Expo.sure ro heat(orradiatiOfl)causes atoms to vibrate violently, and this disrupts hydrogen
and ionic bonds. Under these o:inditions, protein molecules become elongated, disorganised
strantls. We see this when a hen"s egg is cooked. The translucent egg White" is a globular protein,
albumen, which becomes irreversibly opaque and insoluble. Exposure to heavy metal ions, to
organic solvents or to extremes of acidity or alkalinity will all trigger irreversible denaruratiOfls.
These conditiOfls sometime alter charges on the R-groups, too.
Small changes in the pH of the medium may alter the ionic charges of acidic and basic groups
and temporarily cause change (see page 65). However, the tertiary structure may spontaneously
reforrn. This suggests that it is the primary structure of a protein that dcterrnines cross-linking and
thetertiarystructure, givenafavourablemedium.
50
2.3Proteinsandwater .
Roles of proteins
We have seen that some proieins of organisms have a stn.,ctural role in cells and organisms:
OOlers have bioch c mkal or physiological mks. Whatever their roles are in metabolism.
however, cdl proieins are in a continuous state of Oux. They are continuously built up, used and
broken down again into their constituent amino adds, to be rebuilt or replaced by fresh proteins,
accordingtotheneedsofthecells(Figure2.26).
Figure2.26Thedynamicstateofcellproteins
Water
Livingthingsaretypicallysolid,substantialobjects.yetwaterformsthebulkoftheirsaucrures.
Between 65 and 95 per cent by mass d most multicellular plane., and animals (about 80 per cent of a
human cell) consi.sls of water. Despite this, water is a substance that is often taken for granted.
Water is composed of atoms d the elements hydrogen and oxygen. One atom of oxygen and
two atoms of hydrogen combine by sharing d electrons (co,·akm bondlngj. However, the water
moleculeislriangu/arratherthanlinearandthenucleusoftheoxygenatomdrawselectrons
(which are negativdy charged) away from the hydrogen nuclei (which are positively charged) -
with an interesting consequence (Figure 2.27). Although overall the water molecule is electrically
neutral, there is a nei negative charge on the oxygen atom (conventionally represemed by Greek
letter delta as 6 - ) and a nei positive charge on the hydrogen atoms (represented by 6+). Jn Ol:her
12 Distinguish between words, the water molecule carries an unequal distribution of electrical charge within it. This
ionic and covalent
unequal distribution of charge is called a dipole . Molecules which conrnin group, with dipoles are
bonding
known as polar m olecules. Water is a polar molecule.
Hydrogen bonds
With water molecules, the positively charged hydrogen atoms of one molecule are attracted to
the negatively charged oxygen atoms of nearby water molecules by forces called hydrogen
bond'>. These are weak lxmds compared to covalent bonds. yet they are strong e1X>ugh to hold
water molecules together. Hydrogen lxmds largely account for the unique properties of water. We
examine these properties next.
Meanwhile we can note that dipoles are f01Jnd in many different molecules. especially where
there occurs an -OH. -C = O or >N- H group. Hydrogen lxmds can form between all these groups
and also between these group,; and ,vater. Materials with an affinity for water are described as
hydophilic (meaning 'water-loving'. see below).
51
El Biological molecules
Figure2.27Thewater Oll<'""Y9"1\atomcomb<""' umha,edelectrol\S
molernleandtheh)<lrogen withtwoh)<lrogenatom,by (negative charge)
bond,rtforms
~';9elect
j rol\S(covalent ~
"~'"'"' {ff --- : ·-'o)
" " ' ' "<)-
'~ +~'I
, ~ \ '"''"''""00'
=.:::;.:."':.:the ,' -q\-)<? ~:< I -,,
;~r:»::=the ( c;,/x, ~ -- , ~nucleusofh)Orogen
hy<lrngennudeus(positively ',.,,,P ';-' - - , (tl, ,' ;:'!'.';..tor,~)
cha rged) l / / / ~ - - - - ' ' a l \ C jle a ~ - - - - ~ > , ,
thewatermolernlecanies ~ "-
an uneq ual di stribution o f
e lectrica l cha rge.even
though overal l it is
~:.,-&6
electrically neutral triangu la,in,hape.r,otlinea,
j
po la r war rmolecule
there is electrostatic
~:·- · ,,,_dJ
~boodA,.,.,,.
att,actionbetweenthe
pos.itivelycharged,PCJK>nof
Ol\<'Waler moleculea!ldthe
negativelychargedrPCJKlnof
aneighbouringone.gMI\Cj
fisetoweakbond,called ~00000
hydrogen bonds
52
2.3Proteinsandwater .
Consequendy, the evaporation of water in sweat on the skin or in transpiration from green leaves causes
marked ccoling. The escaping molecules take a lOl d energy with them You experience this when you
stand in a draught after a shower. And, since a great deal d he-at is 106! wi!h the evaporation of a smaU
amount of water, cooling by evaporation of water is ==·cat on water oco.
A l>quid at ,oom temperalli!l', water L>qukl medium for living things and !Of the chemist,y
13 Waterhasarelative di,,;ol,esmores.ubstanc:esthanany of life
molecular mass of only othe, common liquid
18yetitisaliquidat
Much heat energy is needed to ,aise Aquatic: environments are slow to cha<>ge temperaru,e
room temperature .
This contrasts with thetemperatureofwate,. Bulkymganismshavestabletemperatu,es
othermiallmolecules Evaporationrequi,esagreatdeal Evaporationcau...,markedcOO;ng.Muchheatislostby
whicharegases. (For ofhe~t theevapo,ation ofasmallquantityofwater.
example.methane Water molecules adhe"' to surface, Water adheres to the wall, of xylem vessels as it is dra,m
{CH,Jwitharelative Wate,columndoesnotbreakOfpull upthestemtotheleavesfrom themots.Watercanbe
molecularmassof16; apartundertension(seeTopic:7 ) liftedbyforcesappliedatthetop,andsocanbedrawnup
ammonia (NHJ with
··-
thexylemvesse!sof~tfeetrunkbyforcesgeoeratedinthe
a relative molecular
mass of 17andcarbon
dioxide{CO,)witha
relative molecular mass
of44.)Whatproperty
of water molecules
mayac:countforthis7
53
El Biological molecules
• Testscan~caniedouttoidentify reducingsugars, non- with ester bonds. The fatty acids maybe saturated or
reducingsuga rs,starch and prote ins. The~nedict'stest unsaturated (containing one or more double bonds between
can be modified to estimate the concentration of reducing carbonatorm in the hydrocarbon tails}. Fats and oils are
effectively energy stores
• Organ ic compounds contain the elements carbon and • ln phospho lipid s,oneofthehydroxylgroupsofglycerol
hydrogen ,u1Uallywithoxygenandotherelements.Carbon reactswith phosphoricacid .Theproducthasahydrocarbon
atoms form 5\rong, cova lent bonds {sharing electrons 'tail' (hydrophobi c)andanionisedphosphategroup
~tweer\ atoms} with other carbon atoms and with atoms of {hyd roph ili c) so they form bilayers on water and form the
other elements, forming a huge range of compounds. Many membranesofcells (alongwithproteins)
oftheor!Jilniccompoundsoflifefallintooneofthefour • Proteimcontain nitrogenand(often)sulfur,inadditionto
group1ofcompounds,ca rbohydrates , lipids, prote in s or carbon,hydrogenandoxygen.Thebuildingblodsofproteins
nucleic acids area minoacids,andmanyhundredsorthousand1ofamino
• Carbohydrates contain carbon, hydrogen and oxygen only, acidresiduesformatypicalprotein.Aminoacidshavea basic
andhavethegeneralformulaofC,(H 10),.Theycomistof am in o group (-NH 2) and an acidic carboxyl group
thesu!Jilrs( monosaccharidesandd isaccharides )and (-COC H) attached to a carbon atom to which the rest of
macromolecules built from 1Ugars {polysacchar ides ) themolecule(-R) isalsoattac:hed
• Monosaa:haridesindudethesix-carbonsugars(hexos es), • Ofthemanyaminoac:idsthatoccur,only20arebuiltupto
1uchasglucoseandfructose.Theyareimportantenergy make the proteins of living things. Amino acids combine by
murresforcells.Thehexoseshavethesamemolecular peptide bonds ~tween the carboxyl group of one molecule
formula (C 6H1iOiJ but different structural formu lae. Two and the amino group of the other to form a po lypeptide
hexose monosaccharides combine together with the removal cha in .Proteim,too,arepolymers
of water (a condensation reaction} to form a disaccharide • Thepropertiesofpolypeptidesandproteimaredeterminecl
sugar, heldtogetherbyag lycos id icbond.Forexample,the bytheaminoac:idstheyarebuiltfromandthesequence
disaccharidesucroseisformedfromglucoseandfructo1e. in wh ich the am in o adds occu r. The sequence of amino
• Monosaa:harides {and some disac:charide1) are reducing acidsisthep rim arystru cture oftheproteinandisindirectly
sugars.Whenheatedwithalkalinecopper{ll)sulfate controlledbythenucleu:s
{Benedict'ssoluti on},whichisblue,theyproducereduced • Thes hapetheprotein molecule takesupalsodeterminesit1
copper{l)oxide,whichisab rickredprecipitate.Therngaris properties.Partofapolypeptidechainmayformintoahelix
oxidisedtoasugaracid.Sucroseisnotareducingrngar. andpartintosheets{theseco ndarystru cture ofaprotein}
• Most polysaccharidesarebuiltfromglucmeunitscondensed Thewholechainisheldinpositionbybonds~tweeriparu
together.Theyareexamplesof polymers consi51ingofahuge folded together, forming the tertiary structure of a protein.
num~r of monome rs, such as glucose, condemed together. Separate polypeptide chains may combine, sometimes with
They may~ food stores {e.g. starch, glycogen) or structural otherrnbstances, toformtheq uaternarystructure
components of organisms (e.g. cellulose, chitin}. • 5orneproteinshaveastructural ro le and are fibrous
• Lipid scontaintheelementscarbon,hydrogenandoxygenbut proteins,forexample,collagenofskin,boneandtendon
the proportionofoxygenis low. They are hydrophobic, Many proteins are enzymes or antibodies {and some are
non polar and insoluble in water but othen\'ise form a hormones}andtheseareg lobu larproteim
diversegroupofcompounds,includingthe fatsandoils • Wateriselectricallyneutralbut,becauseofitsshape,itcarries
The1elattercompoundsleaveacloudywhitesuspensionin anunequaldistributionofcharge-itisa polarmolecule
theemulsiontest. {oxygen with a small negative charge, hydrogen 1\'ith a small
• Fats are §Oiids at room temperature; oils are liquids. They are positive charge). Consequently, water forms hydrogen
formedbyaco ndensationreaction betweenthehydroxyl bondswithotherwatermolecules.Thesehydrogenbondsare
groupsofglyceroland threefattyacids,formingatr ig lyceride, responsible for the properties of water im portant to life
54
Examination style questions I
Examination style questions
1 Pot-,,5aa:harides.suchasglycogen,amylopectinandamylose, d By means of fully annotated skeletal formulae of paru
areformedbypolymerisationofglucose.Fig.1.1~part ofadjacent<:ellulosemolK\Jles,eJq>lainhONcellulose
ofa glycogen molecule. is formed from its component monosaccharide. Explain
how this molec\Jlar strvcturt is tht basis of tht distnctivt
proptrtyofcellulost. [6]
d) De5Cribe tht steps to the test you would need to show
\mtther the monosacchilride formed by hydrolysis was a
reducingsugar. lM\atresuhwouldyouexptct1 13]
e) \Vhefed~collagenoccurinththumanbody1 Ill
f) By means of fully arnotated sk.eletal formulae of partli
ofadjacentcollagenmolt<:ules,explainhowcol!agen is
formed from its component monomtn, and the basis of
thedistinctivepropertyofcollagen [6]
[Total : 20]
3 a) Theanglebetweentheoxygeo-hydrogenbondsinwater
Flg.1 .1 is about 150".8ymeansofafullyannotateddiagram
only,explainwhythetxistenctofthisanglecause5
a) WithreferencetoFig. 1.1, thewatermoleculetobepolar(althoughoverallitis
i) describehowthestructureofglycogendiffersfrom electrically neutral).
thestructureofamylose. [2) b) Outline four oftheunusualpropertiesofwaterthat
ii) describetheadval"ltagesfororganismsinstoring canbeascribedtotheeffectofhydrogenbondsand
polysaccharides, such as gl:,<ogen, rather than stOITig Sl.nlmariwthesi9nificanctofeachforlivin9things.
glucose. 13}
b) Glycogen may be broken OOMl to form glucose 4 a) Chemists often represent the structure of organic
Fig. 1.2 shows r~ion X from the glya:,gen molKU!e in molecules as skeletal formulae in which the C and H
Fig. l.linmoredetail. atoms are left out. Write the sk.tletal formula, for two
molecules of amino add and show the rtaction and
50lutionandonewith2cm1 ofadilutestarchkllutioo,
H OH OH H
wasadded2cm1 ofadilutesodiumhydroxidesolution
Flg. 1.2 andthecontentsgentlyshaken. Thenadilutecopper(ll)
sulfate50lutionwasaddedto tach, drop by drop. The
Draw an annotated diagram to explain how a glucose tubeswef'egentlyshakenbetweenadditions,andthis
moleculeisfOITT1edfromthefreeendoftheglycogen procedurewascontinueduntiladefinitecolourcould
moleculesho,yninFig.1.2 [3] be seen. What colourswooldhaveappearedin the two
[Total: 8] tubes,andwhatdotheresultsindicate? [4]
(Cambridge lntemati0t1al AS and A Level Biology 9700, c) Explainconciselywhatismeantbythefollcming
Paper02 2"" variant Ql November 2008) st ructures of a protein, such as haemoglobin.
i) Primary (2)
ii) Secondary (2)
2 a) Cellulose and collagen are macromolecules. What do you
iii) Tertiary (2)
understandbytheterm·macrornolecule'? [2)
d) lnadditiontohydrogenbonds,threeothertyptsofbond
b) Whencelluloseandcollagenarehydrolysedbyappropriate
areimportantinthesecondaryandtertiarystructure ofa
enzymes,whatarelheproductsineach<:a5e7 [2]
protein. Name and ex.plain the chemical nature of the.e
additional bonds. (6)
[Total: 20]
SS
AS Level
3 Enzymes
Enzymes are essential for life to ex ist. The ir mode of action a nd Th e re are many opportunities in this topic for students to g a in ex-
the factors tha t a ffect the ir activity ue explored in this topic. Prior pe rie nceofca rryingoutpractica linvestigations a ndana lysingand
knowledge forthi s topicis a nunderstandingthat a n e nzyme isa interpreting their results.
biologica l catalystthatincreases th e rateof a reaction a ndre ma ins
unch angedwhe nthereactioniscomplete.
56
3.1 Mode of action of enzymes I
,ymheskofromplex Enzymes as globular proteins
molecu~usedingrCJMh
andde,,..lopmeo! and in Jn Topic 2 we saw that the teniary scructure of glo buL-.r proldns was typically spherical. In these
metabolkprocesses, moleculestheirline:irchainofarninoacids(primarystrucrure)ispreciselyfoldedandheldina
e.9.proteios,
~d!arides, globular three-dimensional shape containing a -helices and J!-sheeis. Also, the R-groups d the amino
lipid,,OO!mooes,growth adds present on !he exterior of the molecule are hydrophilic groups. making the protein water
lactors,haemoglob<n, soluble. Remember, this structure contrasted with that of the fibrous proreiru, such as o:illagen.
""''''··=1
~·
e!l<'fg)'-rek>ilSOng
,eactioos,
1.e;...ction,·c
them. lbe molecules mUSI collide with each other in the right way and at the right speed. If the
an gleofcolHsionisnOlcorrect,themoleculesbounceapart. Alternatively,ifthespeedofthe
colHsion speed is wrong or the impact is too gentle. for example, then there will be insufficient
release of simple energy for the rearrangement of electron,. Only if the molecules are lined up and collide with the
sub'ita»ces, correct energies does a reaction occur.
e.g.smal l ioorganic
The 'right" conditions happen so rarely that the reaction doesn~ happen to a significant extent
~u;:'!:.1~/1>0, normally. If we introduce extreme conditions. such as those lisied above, ,ve can cause the reaction
Figure3.1 Metabol~m:an to happen. On the other hand, if we introduce an enzyme for this particular reaction then the
reaction occurs at great speed. Enzymes are amazing molecules in this respect.
57
El Enzymes
g lucose
whensucroseandwatercollkleat
the wrong speed
Theseeventsarewhathappensat
most,andomccllisions
Forthereactiootooccu,,,ucmse
andwatermustcollidein
justthe,ightway - glucoseand
fructosea,efom,ed
Unde,n0<malrnr>ditionsthis
happens so very infrequently
itisaninsignifKantevent
31nthep,esenceofonemcleruleoftheenzymesucrase(inve,tase),
a,,proximately3.0x104mokecutesof,uaosearehy<lrolywdeachminutel
Figu,..,).2Canareactionoccu,withoutanenzyme?
58
3.1 Mode of action of enzymes I
' boulde ronhillside' mod e lof activation e ne rgy withoutacatalyst,this
amoontofer,ergyl>l'ed,
:~=\ltfr,tostart the effect of catalyst
freeeoergychar,ge
products
Figure3.3Activabooene,gy
Thesequer.c:eofstepstoanenzyme-c..talysedreaction EnzymesaretypicaUylarge
e nzyme+substrate -----E-Scjmplex _____.p,odu ct+enzymeava ilab le for reuse globuL-.rprotcin molecules.
Mostsubsiratemolecules
{sub strate raised to
arequiteSlll3Jlmolecules
lfansitk>nrtate) Q . = rnbstratmrnecute-
by =mparison. Even when
;~::~~0>~~~~)
the.suhstrntemoleculesare
verylarge,suchascertain
u~u
macromolecules like the
polysaccharides,onlyOflebond
inthesubstrateisincontaawith
theacrivesiteoftheenzyme.
Figure3.4Thelod:andkeyhy;,othesisofenzymeaction
59
El Enzymes
This means that an enzyme "reo:ignise:s' a very small group of substrate molecules or even only
a single type of molecule. This is because the active site where the substrate molecule binds ha,
a precise s hape and Jistinc1frc c h c mkal properlics (meaning the presence of panicular
chemical groups and bonds). Only pankular substrate molecules can fit to a panicular active site.
Allothersub.stratemoleculesareunablerofitandsocannotbind.
o!heraminoac:Kl
,esid,,._.,a,..·reac:tive".
:~~=:~
Specificity:
~":9r!.=·
of product(,).
• Someamiooacidre,id,,..,ailcJNa e.g.20aod41
panic:ula,substratemoleculero'fir
• Someamiooacidresid=bringaboot
panicula,d,emkalchaoge,
60
3.1 Mode of action of enzymes I
hnokinas•
(C""V'ler-gene<ated
molecular model)
Figure3.6Compute,-generatedimageofthe;ndun,dfithypothesisinacti0<1
Naming enzymes
Manyenzymeshaveanamebasedonthenameoftheirsubstrate.withtheending -aseadded.For
example, lactasehydrolyseslacroseandamylasehydrolysesamylose.
Other enzymes have been given name., that tell us little or noihing about what they do, such as
many of the enzymes of digeS1i0fl, for example, pc.•p,;.ln, lrypsln and rennin.
l a Oef1netheterm Today, systematic naming of enzymes is based on an agreed classification of enzymes and on
'catalyst'
the name of the substrate catalysed. lbese types of names are long a nd detailed. They are outside
b list two difference,;
the scope of this book. 1bey are used in the communications of enzymologists but not in everycfay
between inorganic
catalysts and usage. However, you are already familiar with certain enzymes. For a:ample, the enzymes that
catalyse the formation of two products from a larger substrate molecule by a hydrolysis reaction a re
clas.sified as'hydrolases'.Canyounameahydrolase?
61
El Enzymes
Measuring the rate of reaction
Inpracticalterrns, therateofanenzyme-caclysedreactioni:sckenastheamountofsubotratethat
has di.s3ppe-ared from a re-action mixture in a given time. Altemati\'ely, the 3mount of product !hat has
accumubted in 3 period of time can be measu=I. For example, in Figure 2.38, the enzyme amylase i.,
usedrodigescstarchrosugar,andhereiti'ltheratethatthesubstratestarehdisappe-arsfromare-action
Working with catalase, however, it is e-asiesc ro measure the rate at which the product (oxygen)
accumulates. In the experiment iJJustrated in Figure J.8. the volume of oxygen that has accumulated
athalfminuteintervalsisrecordedonthegraph.
Jn both of these examples we find that, over a period of time. the initial rate of reaction is not
maintained but, rather, falls off quite sharply. This is typical of enzyme actions scudied outside their
locationinthecell.Canyouthinkofreasonswhy1
Thetesttubeis tipped up
tomixtheenzymesolutKln oxygen produced.
withtl>esuhstrate rnUectedbydownward
~·- \
disp!.Kementofwater
in an inverted measuring
~i
hy<J,oger,
pernxide
solubon
(10volume)
The,ate olanenzyme
reactionisg,eatestatthestart,
the initi a l rate
J
~~5 20
,s ~~I ,a<e , ,,mo, :'.:;~;'.;.~;,,
: ~2
i1
s~ 1s 1: ::
] "10 :: ~!
( S = N
frnm 240 25
the 270 255
graph till\lYs 360 ]00 26
Figun,3.8Measuringtherateol
L If the in;tial rate ofO, production continued lo, 120s. then 28 cm'ofO, would be produced
The,efore theinitial,ate•28/120cm 1, - 1
,eactioousingcata lase •0.23cm1,-•
62
3.2Factorsthataffectenzymeaction .
The fall-0ff can be due to a number of reasons . Most commonly it is because the concentration of
the suhstrate in the reaction mixture has fallen. Consequently. it is the initial rate of reaction that is
measured. This is the slope of the tangent to the curve in the initial siage of reaction . How this is
cakulatedisshowninFigure3.8.
Temperature
Examinetheinvestigationoftheeffectoftemperarureonthehydrolysisofstarchbytheenzyme
amylase shown in Figure J.9. When stan:h is hydrolysed by the enzyme amylase, the product is
maltose. a disaccharide. Siarch gives a blue-black colour when mixed with iodine solution (iodine
in potassium iodide solution) but maltose gives a red colour. The first step in this experiment is to
bring samples of the enzyme and the substrate (the starch solution) to the temperature of the water
bathbeforebeingmixed - astepcalled'pre-incubation'.
63
El Enzymes
Theexpelimentisrepeatedatarangeoftemperatures,,uchasat10,20,30,40,50aml60°C
Othe,variables - sucha,theconcentrationsoftheenzymear>dsubstrate
soltrtkms - werekeptconstant
The progress of the hydrolysis reaaion is then followed
Uptoabout40"Ctherateincreases - a10"Crise
bytakingsamplesofadropdthemixrureon theend
intemperatu,eisaccompan>edbyanapproximate
doublingofthe,ateofreact,00 of the glass rod, at half minute intervals. These are tested
withiodinesolutiononawhitetile. Initially. a strong
~.t ( blue----blackcolourisseenconfirmingthepresenceof
starch. Later, as maltose accumulates, a red colour forms
Y\
The end point d the reaction is when all the starch
colourhasdisappe-aredfromthetestspot.
,,;'.::~7",.:. s.
denaturatoonof
temperatures, say at 10, 20. 30, 40, SO and 6()0 C. The
time taken for complete hydrolysis at each temperature
theenzymear>d isrecordedandtheratedhydrolysisinu n ittimeis
destructKlnof plottedonagraph. Acharacteristiccurveistheresult -
a,ct;...,,rtes
although the 'optimum' temperature varies from reaaion
to reaaion and with different enzymes (Figure 3.10).
Hou1istbe graphinterpretcd?LookatFigure 3. 10.
10 20 30 40 50 60
temperature at whoch the ,ate was measure<V'C
t \
w
enzymeinactivestate - ~ =:;i;te:~"f!c'::,~
nolongerfitthe.tctivesite
= C)
Figure 3.10The effect of temperature ootherate of an enzyme-catalysed reaction
64
3.2Factorsthataffectenzymeaction .
4 lnstudiesoftheeffect As the temperature is increased, molecules have more energy and reaction., between them h3ppen
of temperature on more quickly. lbe enzyme molecules are moving more rapidly and are more likely to coUide and
enzyme-catalysed =cr. In chemical reactions, for every !(fC rise in temperature the rate of the reacrion approxinl3tdy
reaction1,suggest doubles. This property is known as the 1cmpcr.11urc coefficient (Q,o) of 3 chemical reaction.
whytheenzymeand However, in enzyme--carnlysed reaction., the effecr of tempernture is more compl= because
s.ubstrate'iOlutK>ns proteins 3re d c natun.-d b y h ea l. The rate of denaturation increases at higher temperatures, too. So
arepre-inrubatedtoa as the temperature rises above a cerrnin point the 3mount d acrive enzyme progressively decreases
partirnlartemperature
and the rate is slowed. As a result of these two effects of heat on enzyme-<:arnlysed reactions, there
before they are mixed
isanapp3rentop1imumtemperatureforanenzyme.
NOi all enzymes have the S3nte optimum temperature. For example, the bacteria in hot thermal
springs have enzymes with optimum temperatures between 80 and I00°C or higher, whereas
seaw=ds of northern seas and the pbnts of the tundra have optimum temperatures closer to 0°C.
Humans have enzymes with optimum temperatures at or about nom,3l body temperature. This
feature of enzymes is oflen =plaited in the commercial and industri3l uses of enzymes today.
pH
Change in pH can have a dramatic effect on the rate of
an enzyme-catalysed reaction. Each enzyme has a range
of pH in which it functionsefficiendy - oftenatorclose
to neutrality. The pH affects the rate of reaction because
the structure of a protein (and therefore the shape of the
active site) is maintained by various bonds within the
three-dimensional strucrure of the protein (Figure 2.22). A
change in pH from the optimum V3lue alters the bonding
patterns. As a result, the shape of the enzyme molecule
is progressively changed. Theactivesitemayquickly
become inactive. However, the effects of pH on the active
sitearenormallyreversible(unliketempcraturechanges).
lowerpH optimumpHforenzyme highe, pH
That is, provided the change in surrounding acidity or
I I I alkalinity is not too extreme . As the p H is brought back to
s.ubstfatemolecules enzyme on substfatemok>cules the optimum for that enzyme, the 3ctive site may reappe3r
oolonge,fitthe active state oolooge,fitthe
act~srte i iK!i~srte
(FigureJ . ll).
Someofthedigestiveenzpnesoftheguthavedifferen!
' (aj ' optimum pH V3lues from the majority of other enzymes. For
~ ~ CiJ
=ample, those adapted to work in the stomach, where there
isa high concentration of acid during digestion, have an
optimum pH which is dose to p H 2.0 (Figure J.12)
structureofproteinchaogeswhenac:hangeofpHalters the
ionk charge on -COO\;,cidk) and - NH, *{basic} groups in the
peptidec:hain,sotheshapeoftheiK!M,siteislost
Figurel.11TheeffectofpHonenzymeshapeandactMty
5 a Explainwhata
buffer'iOlutionis
b Whyaretheyoften
used in enzyme
experiments?
Figurel.12Theoptimumpl-1of
""'
different human enzymes
6S
El Enzymes
Substrate concentration
The effect of differern concentrations of substrate on the rate of an enzyme-catalysed re-action can
beshownusingtheenzymecatalase.
Look again at the investigation of the initial rate of reaction using catalyse (Figure 3.8). We saw
that, when working with catalase , it is easy ro measure the rate the product (oxygen) accumulated
(recordedathalfminuteintervals).
To investigate the effect of substrate concentrntion on the rate of this enzyme-catalysed reaction, the
experimern shown in Figure 3.8 is repeated :ll diffcrenl conccnlrntions of s ubstrate. lbc initial rate
of reaction planed in each case. Other variable., such a., temperature and enzyme coocentration are
kept constant.
When the initial rates of reaction are planed against the substrate CClflcentration, the curve
shows two phases. At lower concentrations the rate incre-ase:s in direa proportion to the substrate
concernration but at higher substrate concentrations. the rate of reaction becomes constant and
6 Whenthereisan showsnoincrea.sc(Figure3.13)
excess of substrate We can see that the enzyme catalase does WOfk by forming a shon-lived enzyme-substrate
presentinanenzyme- compl=. Al low concentrations of substrate, all molecules can find an active site without delay.
catalysedreaction, Effectively. there is =cess enzyme present. The rate of reaction is set by how much substrate is
explain the effects on
present - a.smoresubstrateismadcavanabletherateofreactionincreases.
the rate of reaction
ofincreas.ingthe At higher substrate concentrations there are more substrate molecules than enzyme molecules.
concentration of· Now , in effect. substrate molecules have to 'queue up' fOf access to an active site. Adding more
a thesub1trate substrate increases the number of molecules awaiting contact with an enzyme molecule. There is
now no increase in the rate of reaction. This relationship between amount of substrate and the rate
b theenzyme.
of reaction is shown in Figure 3.13.
0 0(9 O
6J O U\v
00
\ __,,0
o 6Jo 6J
law substrate molecul es. ______. more substrate molecules. ______. H<ffs of substrate molecules.
=~:.!:::conm,1ra1K>ll
wilinaease therate
~~m~::'te~;,catalysis ::::;::.i;,,~!::a~
wil ootcllangethe,ate
hi!f, t t
~i
high
Figure 3.13Theeffectofsubstrateconcentratiooontherateofanenzyme-catalysedreaction
66
3.2Factorsthataffectenzymeaction .
Enzyme concentration
If there are plenty of substrate molecules in a reaction mixture, then the more enzyme that is
added the fascer the rate of reaction wUJ be. This is the situation in a cell where an enzyme reaction
occurs with a small amount of enzyme present. Any increase in enzyme producrion will lead to an
increased rate of reaction. simply because more active sites are made available.
-.L:-r
concemration, the velocity increases - rapidly at lower substrate concentrations. However, the rate
increaseprogressivelyslows,andaboveacertainsubstrateconcentration,thecurvehasOatrened
out. No further increase in rate occurs. This rells us the enzyme is working at maximum velocity at
this point . On the graph, this poim of maximum velocity is shown as v .....
~ l =x --
i
- I, Km
substratecoocentratkm
Figure3.14Graphofinitial,ateclanenzyme-catalysedreactiooagaimtsubstraternncentrahon
lnl913thebiochemistsMichaeHsandMentensrudiedthisaspectofenzpnereactions
and introduced a COflstant, now known as the Michaelis-Menten consmnt. The Mich aelis-.
Mc nte n con s ta nt (K..)isdefinedasthesub.strateconcentrationthatsustainshalfmaximum
velocity <1V...J. h measures the degree of attraction or affinity of an enzyme for the substrate - the
smaller the K,. the higher the affinity. K,. can be =perimentally determined at a specified pH and
temperature,andis=pressedinunitsofmolarity.
Actual values of K,. have been measured for a great many enzymes. These values fall between
10--1 and 10...,, moldml of substrate. This is a very wide range of concentratiOfls. It means that some
enzymes are able to work at maximum velocity at very low concentrations of substrate, whilsc
cxhersonlyfunctioneffectivelyatmuchhigherconcentratioru.
Ta ke the case of two enzymes that camlyse the transformation of the same substrate molecule
but in different re3cti0fl sequences. If the 'pool' or reserves of that substrate are low and its supply
restricted, then the enzyme with the lowest K., will claim more - and one particular metabolic
pathway will benefit at the expense of the other. Knowing the K., of enzymes is an important part
of understanding metabolism, both quantitatively and qualitatively.
67
El Enzymes
Inhibitors of enzymes
Certain substances present in cells (and some which enter from the environment) may reaa with
an enzyme, altering the rate of reaaion. These suhsrnnces are known as inhibitors, since their
effectisgenerallyrolower therate ofreaction.Siudiesoftheeffectsofinhibitorshavehelpedour
understanding of:
• the chemistry of the active site of enzymes
• the natural regulation of metabolism and which pathways operate
• the ways certain commercial pesticides and many drugs work (by inhibiting specific enzymes and
prcventingpanicularreaaions)
For example, molecules that sufficiently resemble the submrme in shape may compete to occupy
the aaive site. Th'-;' are known as compctith-c inhlbilors. For example, !he enzyme that catalyses
the reaaion between carbon dioxide and !he CO,-accepior molecule in photosynthesis, known as
ribulosebispho.sphatecaJboxylase(rubisco),isccrnpetitivelyinhibitedbyoxygeninthechloroplasts.
Because these inhibitors are not acted on by the enzyme and turned imo 'produas' as normal
substrate molecular are, they tend to remain attached. However, if the concentration of the
suhstratemoleculeisraised.theinhibitormoleculesa1tprogressivelydisplacedfromtheactive
sites(FigureJ.15)
Alternatively, an inhibitor may be unlike the substrate molecule, yet still combine with the
enzyme. In these cases, the anachment occurs at some other part of the enzyme. This may be
quite dose to the active site. Here the inhibitor either partly blocks access to the active site by
substratemolerulesoritcausestheactivesitetochangeshapesothatitisthenunabletoaccept
the substrate.
l1>ese are called non.competitive inhibitor,,, since they do not compete for the acrive site. Adding
o:ccsssubstratedoesnot=-theirinhibitingeffa:ts(Figw-eJ.15).Anexampleistheeffectofthe
amino acid alanine- on the enzyme P')-nJvate kinase in the final step of glycolysis (Topic 12)
olenzyme-cataly,,<lre,o::!ion
- ~~7: ~,a 1
competitive
substrateaodinhibilo,
competefo,acti,,..,ites,so
I ,;--c:·.,::,,oooom
.,o,.·,-,
V ~~~1:~l~
1
68
3.2Factorsthataffectenzymeaction .
Certain irreversible inhibito r s bind tightly and permanently to an enzyme and destroy its catalytic
properties entirely. These drastic effects occur at low concentrations of inhibitor and we may
describe these subsrnnces as poison s. Examples include:
• cyanide ions which block cytochrome oxidase in terminal oxidation in cell aerobic respiration
• thenervegassarinblocksaneurotransminer(acetylcholinesterase)insynapsetransmission
Application
'"""" 'BK>logical"detergents
fructa5esyrupmanufacture
Fung al Kluyveromyces BreakdownoflactosetoglucoseaOOgalactme
Amylase Removalofstln:hinwovendothpmduction
69
El Enzymes
Immobilised enzyme - a laboratory demonstration
The steps to this demonstration are
I A solution of sodium alginate (a polysaccharide obtained from the walls of brown algae,
capable o f holding 200 times its own mass in water) is prepared by dispersing 2g of the alginate
inl 00cm 1 ofdistUledwateratatemperarured40"C.
2 An alginate-enzyme solution is prepared by stirring 2cm1 of invertase concentrate into 40cm1 of
thecooledalginatesolution.(Remember, invertasecatalysesthehydrolysisdsucrose[a
d isaccharidel to two monosaccharide molecules (glucose and fructose) from which it is formed
inacondensationreaaion.)
., lmmobm.sed enzyme pellets are produced by filling a syringe with the alginate-invertase
solution and arranging for it to drip into a beaker of calcium chloride solution (IOOcm 1 o f CaCl,,
O.lmolilllll). The insoluble pellets formed may be separated with a nylon/plastic sieve. They
shouldbewashedwithdistilledwateratthisstage,andallowedroharden
4 lbese hardened pellets may be tested by placing them in a cube with a narrow nozzle at the base
(the barrel of a large , plastic syringe is suitable). A quantity of sucrose solution (JOcm1 of 5%
sucrose w/v) may be slowly poured down the column of pellets and the effiuent colleaed. Note
thetimetakenforthesolutiontopas.sthrough
5 Te~t the effluent solution for the presence of glucose, using dinistix™ (see below).
Preparing immobilise d e nzyme in a lgin at es Harvesting a lginate-e nzyme beads Using immobilised e nzyme beads
s.ubstrate 5rnution
tricklesslo\\iypast
the beads
products of the
enzyme reaction
(glucose+fructosesolutoon)
Figurel.16Prepa,ir,ganimmobilisedenzyme
70
3.2Factorsthataffectenzymeaction .
the principles
glucoseoxidase
gltJCose+oxygen ---- gluconic:add+Hz()1
peroxidase
DH1+H10., 2Hz()+O
reduced chromagen
chromagen (coloured)
(coloorle<ss)
Advantages Disadvantages
The enzyme is held in a form that can be manipulated An immobilisat>OI\ mechanism that does not alter the
easily.ltisabsentfrnmtheproduct,sonopufif,cat>OI\ shapeorthecata!yticab<lityoftheenzymemustbe
steps are ,equired se lected
Theenzymeisava ilableformultip lere·use.sinc:eit Thecreationofstable.h,wJenedpelletsisanadded
functionsasaneffectivecatal,,stinpelletfOf!TI expensethatlsinevitablyreflectedinthecostofthe
k,dustfialproduct
An immobilised enzyme is stable at the tempe,aturPS tt the enzyme becomes detached it will appea,in the
andpHatwhichitisheldandused productasacontaminant.possiblyunnohced
71
El Enzymes
loralurthertenminutes
Alter ten minutes, the temperature of the water bath was Fig.1.1
rai1edtoboilingpoint.Benedict'ssolutionwasaddedtoeach
test-tube.Thetimetakenloracolourchangewasrecorded 2 a) Describe how enzymes take part in chemical
andusedtocakulateratesolenzymeactivity. reactions (4)
TheresultsareshwminFig.1.1 Starchphosphorylaseisanenzymefoundinplantcells.ln
a) i) Namethetypeolreactioncatalysedbysucrase [1) potato tuber cells, the enzyme takes part in the breakdown
ii) Explain why the temperature of the water was rai sed olstarchwhenthetuberbeginstogrow.
to boiling point [2) starc:hphosphorylase
b) Describe and explaintheresultsshCM'n in Fig. 1.1 [5) starch+phosphateions--tglucoie-1-phosphate
[Total:8] AstudentinvestigatedtheeflectofpHonthisreactionusing
(Cambridge International AS and A Level Biok;,gj 9700, two buffer solutions
Paper21Q41une2011) The student prepared lour test tubes, A to D, a1 shown in
thetableanddescribedbelow.
Thestudentmadeanextractolpotatotissuethatcontained
the enzyme. Some of this extract was boiled.
72
Examination style questions I
A mlution of pot;m.ium dihydrogen phosphate was added ii ) ExplainwhythestudentOOledsomeoftheextractin
tosometubesasa1ourceofphosphateions. this investigation [2]
Thetesttubeswereleftfortenminutesinawaterbathat c)Explaintheresultsshowninthetable [4]
30°Candthensamplesweretested\\ithiodinesolution. [Tota l: 11]
b) i) State what the student would conclude from a (Cambridge International AS and A Level Biology 9700,
positiveresultwit hiodinesolution (1] Pape< 02 Q2 June 2007)
negative
3 Thediagramshow:sanapparatususedinaninvestigation
using immobilised enzymes. It is no t expected that you will
have done this investigation
Asolutionofasubstratewaspouredintoaburette ,obora• wl•A
containinganenzyme immobilisedonto alginatebeads.The
liquidpassingthroughtheburettewascollectedintoabeaker lm=boB,od=,...lo
andtheconcentrationofsubstrateandthernncentrationof bunma
the product measured. Thetableshowstheresultsobtained
by five students
liquidcollaclad
u - - lromburn"8
73
AS Level
74
4.1 Fluid mosaic membranes I
water upta ke/ loss
neurot.-ammittersubstanc:es
hormones
# ,'o""'") :::;,~;:i:,o;orn
collagen fibres
outside the
mammalian eel)
o,
components -
celluloseand
hemicellukise
(assembled to
make plant cell
wall.outside eel~
ions
e.g.Na•,K•,c,l•,a!so
t,aceelements(e.g.fe,Cu)
Fig ure 4.1 ThemoYementofsubstancesacros,thecellsu,facemembra""
You can see thm the pho.spholipid molecule has a 'head' composed of a glycerol to which is
attached an ionised phosphate group. This latter pan of the molecule has hydrophilic properties
(water-loving). For example, hydroge n bonds readily form between the phosphate head and
water molecules. The remainder of the phospholipid comprises two long, fatty acid residues
consisting of hydrocarbon chains. lbese 'tails' have h}'dro phobic propenics (water-hating).
!!!!!/"""
Alternatively, the phospholipid molecules arrange themselves as a bilayer, again with the
hydrocarbon tails facing together (Figure 4.2). This is how the molecules of phospholipid are
arrangedinthecellsurfacemembrane.
In the phrupholipid bilayer. anractions between the hydrophobic hydrocarbon tails on the inside
When mlndwlthwat.r, andbetweenthehydrophilicglycerol- phosphateheadsandthesurroundingwaterontheout.side
~molec:ule<anange make a stable, strong barrier.
t ~ i n r o a bllayu,in
Finally, the phospholipid bilayer has been found to contain molecules of c holcs 1crol
~.~~~~!,are O>olesterol has an effect on the Oexibility of the cell surface membrane. We return to this fearure
shortly.
75
IJ Cell membranesand transport
The proteins and carbohydrates of membrane
The proteins of membranes are globular pro teins which are buried in and across the lipid bilayer,
with most projecting above the surfaces. O!hers are superficially attached on either surface
o fthelipidbnayer.
Proteinsthatoccurpaniallyorfullyburiedinthelipidbnayeraredescribedasintcgral
proteins . Those that are superficially attached on either surface of the lipid bilayers are known as
pcrlphcral protein....
Membrane proteins have a range of roles. Some are enzymes, o thers are receptors. or antigens,
and many are channels for transport d metabolites. ThO'ie that are involved in transport of
molecules across membranes are in the spoclight in the next section. A summary of the various
roles of membrane proteins is given in Figure 4.24 (page 94). afier the movements of molecules
acrossthemembranehavebeendiscussed.
The carbohydrate molecules d the cell surface membrane are relatively short chain
polysaccharides, some attached to the proteins (glycopro1ei11.~) and some to the lipids
(glycollplds) Glycoprotein.s and glycolipids are only found on the outer surface of the membrane.
Together these form the glycocalyx. lbe roles of this glycocalyx are:
1 lnplantsadaptedto • cell---cell recognition
survive in very low • as receptor sites for chemical signals. such as hormone messengers
winter temperatures, • to assist in the binding together d cells to form tissues.
what seasonal change
would you anticipate
in the composition of The fluid mosaic model of membrane structure
thelipidbilayeroftheir The molecular structure of the cell surface membrane, known as the fluid mosaic model by those
membranes? who first suggested it, is shown in Figure 4.J. lbe membrane they described as fluid because the
2 Whatisthedifference components Oipids and pro1:eins) move around within their layer. In fact the movements of the
between a lipid bilayer
lipid molecules are rapid, whereas mOOne proteins move about more slowly. The word mosaic
andthe'double
membrane' of many described the scattered pattern d the proteins, when viewed from above. This is the current view
organelles? of the structure of the cell surface membrane. It is based on a range of evidence, in pan from
studieswiththeelectronmicroscope(Figure4.4).
76
4.1 Fluid mosaic membranes I
Composition of the membrane and control of how fluid it is
lhe phospholipids of the membrane are a mixrure. Some have sarurated fatty add mils and some
unsaturnted fatty acid tails (Figw-e 2.14, page 41). An excess d unsaturated fatty acid tails makes the
membrane more fluid lnis is because the kinks in the tails prevent dose-packing of the lipids. However,
the presence of cholesterol among the pho,pholipid molecules has to be taken into account. The effect
ofcholesterolistoreducefluiditybypreventingorreducingthemovementsofthelipidmolecules.
Membrane fluidity is an important factor. Membranes must be sufficiently fluid for many of the
proteins present to move about and so to funaion o:irrectly. If the temperature of a membrane
falls it becomes Jess fluid . A point may be reached when the membrane will actually solidify. Some
organismshavebeenfoundtovarythebalancebecweensaturatedandunsaturatedfattyacids and
the amount of cholesterol in their membranes as ambient temperatures changes. Jn this way they
maintain a properly functioning membrane, even at very low temperarure.s, for example.
m~
memb rane lineoffractu,eof
membrane shown
to the right
figure 4 .4 Membtane structure: ev.Jenc:e from Ifie e!eclfO<l mKroscope
77
IJ Cell membranes and transport
The fluid mosaic model Sy the end of this section you should be able to:
allows an understanding of
howsubstan(esenterand a) descr ib e andexplainthe proce sses ofdiflusion,facilitateddiffusion,osmosis,active transport,
exit(ellsbyavarietyof e ndocytosisandexocytosis
different mechanisms b) invertigate simple diffusionusingplanttissue andnon-livingmate rial,,suchasglucosesolution,,
Investigating the effect Visking tubing and agar
ofincreasingthesizeof c) ca kulate surfac:e a,e asandvolumesofsimpl e shapes(e .g.cubes)toillustrate the prindple that
model cells allows an surf e a , eatovolume rat·o,de cre u e w·th"ncre a<ng<ze
understanding of the d) invertigate theeffectofchangingsurfaceareatovolume ra tioondiffusionusingagarblocksof
constraints of obtaining
resources across the
e ) invertigate theeffectsofimme rsingplanttissue sinsolutionsofdiffe rentwaterpotential,using
(ell surface and moving
substances out of cells there sultstoestimate thewaterpotentialofthe tis.sues
f) explain th e movement of water be twee n cells and solutions with diffe re nt water pote ntials and
explainth e differe nt effectsonplantandanimalcells
Into and out of cells passes wa ter, resplr:u ory gase,,i (0 2 and CO,), nutricnl.. , cse-cntial mineral
lon~ and excretory products . Cells lllllY secrete sub.stances such as h o rmones and c n Z)'lll""1.
They may receive growth s ubs ta n ces and h o rmones, too. Plants secrete the chemicals that
lllllke up their walls through their membranes. and assemble and maintain the wall outside the
membrane. Cenain mammalian cells secrete s trucmral p ro!cln s such as collagen in a form that
canbeassembledoutsidethecells.
In addition. the membrane at the cell surface is where the cell is identified by surrounding cells
and organisms. For =ample. protein receptor s ites are recognised by hormones, neurotransmitter
substances (from nerve cells), as well as other chemicals, sent from other cells.
We will now look into the m ech.~nisms o f m e mbran e !ra ns port. Figure 4 .5 is a summary of
the mechanisms of transpon across membranes.
1diffus ion
- ~ntf,omhighto
lowconcentfation;
energy source•kinetK
energyofmolecu1es
3 bulktra nspo rt
tfansportofso1idsand/or
liquidsbyvl'Sicleatthe
cellsurfacememb<ane,
' energy source•enen:Jy
from metabohsm
i
2ac tivetranspo rt
select,,.,mOYementofsubstancPS.
aga inst~coocentfation gradient;
energy source • energy from metabolism
Figure 4.5 Mechanisms of mOYement across memb,ane,
78
4.2 Movement of substances into and out of cells I
Diffus io n:thenet Movement by diffusion
movement of particles The atoms, molecules and ions of liquids 3nd gases undergo continuous random movements. These
!.UChasmolerulesfrom movements resul! in the even distribution of the components of a gas mixture and of the atoms,
aregionwheretheyare molecules and ions in a solution. This is why we are able to rake a tiny random sample from a
atahigherconcentration solutionandanalyseitto findtheconcentrationofdissolvedsubsrancesinthewholesolution.
toaregionwithalower
Every sample haa the same composition as the whole. Similarly, every breath we rake has the same
concentration,using
amount of oxygen, nitrogen and carbon dioxide as the atmosphere as a whole
energy from the random
movements of particles G:intinuous random m - = t s of aU molecules ensures complete mixing and even dismb.nion of
Thi1includesdiffusionof molecules, given time. in solutions and gases. lbe energy for diffusion comes from the kinetic <'1lefl,'Y
small non-polar molecules ofmolecu.les. 'Kinetk'me-:,nsthatapan:iclehasthi'lenergybecausei!i'lincontinuousmaion.
{suchasoxygenandcarbon Diffus io n is the free passage of molecules (and atoms and ions) from a region of their high
dioxide)throughthecell concentration to a region of low concentration. Where a difference in concentration has arisen in a
surface membrane, as well gas or liquid. random movements carry molecules from a region of high concentration to a region
a1diffusionoffat-soluble of low concentration. M a result. the panicles become <,>venly dispersed. The factors affecting the
molecules(suchas rateofdiffusionareli.stedinTable4. l .
vitaminNthroughthecell
surface membrane.
Table4.1Factorsaffectir,gtherateofdiffus""1
Theconditionsneededtoachieverapiddiffusion
Factorsaff.ctingtherateofdiffusion acros1a1urf e
Concentfationg,adient - thegreate,thedifferenc:ein Afreshsupp!yof,ubstanc:eneedstoreachthesurfac:e
c:oncentfationbetweentworegions,thegreate,the aOOthesubstanc:ethatha,c:rossedr,eedstobe
amount that diffuses in a gWen time transported away.
distanc:e.thegreate,the,ateofdiffusion
Area;,c:rosswhic:hdiffu,ionocc:urs - thelarger the Tfle,urfacearear>eed,tobe!a,ge
area.thegreate,thedilfus.ioo
Structurethroughwhkhdiffus.iooocc:u,s - pores0< Ag,eaternumbe,ofporesar><la!arge,sizeofpores
gapsinstroctu,esmayenf>anc:ed ilfusion ,myenhanc:edilfusion
Sizeandtweofdiffusir,gmolecules - ,ma lte, Oxygenmaydiffusemorerapid!ytf>ancarbondioxide
molec:utesandmoleculessolubleinthesubstanc:eofa Fat-solublesubstanc:esdiffusemore rapid!ythrnogh
barfie,wi llbothdilfusemorerapid!y. the lipidbi!aye,thanwater-solublesubstanc:es
Demonstrating diffusion
DiffusioninaHquidcanbeilJustratedbyaddingacrystalofacolouredmineraltodistrnedwater.
Even without stirring, the ions become evenly distributed throughout the water. The process takes
time, especiallyasthesolidhasfirsttodissolve(seeFig ure4 .6 onthenextpage).
79
IJ Cell membranesand transport
1 Aay;talofpot.N;ium 2Astheior,;;di,ssolve, 3Theionsbec:omeevenly
pem>anganate (po tassium -----+ random ma,ements -----+ distribl11ed . Random movements
manganate(VII). KMn0 4) is disperse them through continue. bl11 there is now no net
placed in distilled water the wate, movement in any partku!a, direction
Figure4.6Dif1usioninaliquid
3 Forimaginarycubic
'cells'withsidesl,2,4
and6mm·
a cakulatethevolume,
surface area and
ratio of surface area
to volume for each
b plotagraphofthe
surlaceareaofthese
cells against their
volume
c statetheeffecton
theSA :V ratioofa
cellasitincreasesin
d explaintheeffectof
increasingcell!.ize
on the efficiency
ofdiffu!.ioninthe
removal of waste
surface area
products from cell \OloJmeratio
6:1=6t,=6 24:a=24t8 =3 54:27='>4t.17 =2 96:64=116t64 =1.5
cytopla1m.
Figure4.7Theeffectofinc:,easing,;zeonthesurlacea!l'a :vffiumeraho
80
4.2 Movement of substances into and out of cells I
4 Cubesofslightlyalkalinegelatinofdifferentdimens.ions,containinganacid-alkaliindirator
(redinalkali1butyellowinacids}wereprepared. Thecubes1verethenplacedindiluteacid
solutionandthetimetakenforthecolourinthegelatintochangefromredtoyellowwas
measured
I Dimension s/mm I Surfacean,a/mm• I Vo1urnelrnm 1 I Time/minutes I
l 10x10x10 lsoo 1,000 In I
l sx5x5 l 1so Im 14.S I
l 2.sx2.sx2.s 137.5 1 15.6 140 I
a Foreachbkx:k,calculatetheratioofsurfaceareatovolume{SAIV)
b Explain why the colour changes more quickly in some blocks than others
Diffusion in cells
• the membrane is fully permeable ro the substance concerned. The lipid bilayer of the cell
surface membrane is permeable to non-polar substances, including steroids and glycerol, and also
oxygenandcarbondioxideinsolution,allofwhichdiffusequicklyvfathisroute. Nore that the
ne1 diffusion of different types of panide can mke pbce in opposite directions without hindrance
too. So, at the lung surface. =ygen diffuses into the blood whilst carbon dioxide diffuses out.
• the pores in the membrane are large enough for a substance to pass through. Water diffuses
across the cell surface membrane by means of the prot.ein-lined pores of the membrane. The tiny
spaces between the phospholipid molecules are anOlher route for water molecules. This latter
occurs more easily where the fluid-mosaic membrane contains pho.spholipids with unsaturated
hydrocarbon tails, for here these hydrocarbon tails are spaced more widely. In this state, the
membrane is consequently especially "leaky' to water, for example
po!a,molecules.e.9.wate,.in
molec:ul"".ofaooo-polar,ubstance. 0
Mghrnncentrabc,n ~
t:?<\J~~i~:o~
d-,ffu- ,rn
:::~t;:'o/o
.,.j-. -.- .- .- .-
- be__ - , ,•
o o o o
diffusionoccu"inbothd;rec1'°",.butthere
o
o
O
o
o o
oo
O
o
o
o
o
c
O
o
o
is net movement ffom a region of h>gh net diffusion of polar molecules to a region
coocentrat'°"toa,egionollowcoocentrat'°" 'M\eretheyareinlONerrnncentfation
Figure4.8Drflusionacros,thecellsu,lacememb,ane
Facilitated diffusion
lnfacilimreddiffusion, a substancethatOlherwiseisunablerodiffuseacros.sthecellsurface
membrane does so. This is as a result of its effect on a panicular prot.ein in the membrane. These
are globular prot.eins that can form into pores large enough for diffusion. Those pores dose up
ag:,inwhenthatsubstanceisnolongerpresem(Figure4.9)
81
IJ Cell membranes and transport
Figure4.9FacilitateddiffU1ion
Alternatively a channel prO{ein may be selective. For example, some channel pro1eins are 'gated"
and open to allow the passage of ions only under panicular conditions. There are different
channels for porns.sium ions and sodium ions in the membrane of nerve fibres, for example
(Figure!S.6,page317).
Such prote ins undergo rapid shape changes when in contact with a panicular solute molecule
Facilitated diffusion follows. The movement of glucose into red blood cells occurs in this way. So
does the movement of ADP into mitochondria and the movement of ATP from mitochondria into
5 Distingui1hbetween
thecyrosol.
'diffusion'and
'facilitateddiffmion' Finally, we mUS{ remember that in facilitated diffu.sico the energy comes from the kinetic energy of
themoleculesinvolvcd.asisthecaseinaUformsofdiffusion. Energyfrornmetabolismisnotrequired.
We saw in Topic 2 that in a solution, a dissolved substance anraas polar water molecules around
it (Figure 2.28, page 53). The forces holding those water molecules in this way are h ydrogen
b o nd ... The effect of the presence of a 'cloud" of water molecules around each solute molecule is
to hamper and re.stria their movements. Organic sub.stances like sugars, amino acids, polypeptides
and proteins, and inorganic ions like Na+, K"._ Cl - and NO;- , all have this effect on the water
molecules around them. On the other hand, in pure water. all of the water molecules are free to
move about randomly. and do so - all the time. We say that here they diffuse freely.
Jn the =periment in Figure 4.10. the solution of sugar was separated from pure water by
a membrane - the walls of a dialysis tube. This is described as p.~rtially perm eable because
it only allows cenain molecules to pass, no1 all of them. Water molecules move across the
membrane in both directions, by diffusion. At the same time, the solute mol=ules with their
cloud of water molecules are too large to pass through. In the mean time. there is a continuing
net movement of water mol=ules from the region of high concentration of free water mol=ules
(the molecules of pure water in the beaker) to the region of low concentration of free water
molecules (the water mol=ules of the sucrose solution). This causes the bag to fill up and
become stretched and turgid.
82
4.2 Movement of substances into and out of cells I
, ~'.:;.':""""" \ :::,;:::::".:..,.,
netlklw
of water
intotheba.g
Figure4.10 0;mosis
Water potential
The name given to the tendency of water molecules to move about is wa ter po lc ntia l. 'Water
pocential' is really a measure of the free kinetic energy of the water molecules. The Greek letter psi
(symbol 111) is used to represent water potential.
Water moves from a region of higher water potential to a region of lmver water potential. We
say water moves down a water poten tial gndlcnt . Equilibrium is reached only if or when the
water pocemial is the same in boch regions. At this point, there would be no net movement of
water from one region to anO!her. but random movements d water molecules continue, of course.
Next we will =amine the effects of dissolved solutes, and then of mechanical pressure on water
potential, before returning to a re-statement of water pocential.
83
IJ Cell membranesand transport
Thisexpe,imeotcanbe 1e...,1ofsolutionris'-" The concentration of solute molecules
carriedout inyoorlabofato<y
and water potential
1 Pure water obviously has the highest W3ter potenti3l. By convention.
this is set at zero. Once 3 solute is dissolved in W3ter, the W3ler
molecules are immediately less likely to diffuse (they are less mobile).
So the effect of dissolving solute in water is to lower its water
potential. Consequently solutions 31 3tmospheric pressure must have a
negative value of water potential (since pure water is set at zero). Also,
sucrose solution thestrongerthesolution(i.e. themoresolutedissolvedpervolumeof
(sucrose is the solute) water) the l3rger the number of W3ter molecules that 3re slowed up
and held almost stationery. So,in3veryconcentratedsolution, very
m3ny more of the W3ter molecules have restricted movements than do
in 3dUutesolution.
The amount of dissolved solute present in a solution is known
purewate, as the soluu, pok·nt la l of the solution. It is given the symbol ¥', · A
(water is the solvent) simple o.smometer is shown in Figure 4.11. We use an osmometer
to demonstrate the solute porential of a solution. Once the
o.smometer is lowered into the beaker of W3ter, very many more
W3ter molecules diffuse 3cross the membrnne into the solution
partiallypermeable - ---,----,CLU.LU than move in the opposite direction. The solution is diluted 3nd
membrar,e,e .g
it rises up the attached tube. An o.smometer like this could be
dialysismemb,ane
used to compare the solute potentials of solutions with different
Pressure potential
The other factor that may influence osmo.sis is mechanical pressure acting on the solution. This
factor is given the name p ressure po lcntia l. It is represented by the symbol ¥',,.
Jfapressuregrearerth3n3unosphericpressureisappliedtoasolution,thenthisis3nexample
of a pressure potenti3l being created in 3 solution. Jn the demonstration in Figure 4 .12 a short
length of dialysis tubing has been set up to show how the pressure potential of a solution can
becomelargeenoughtostoptheosmoticuptakeofw3teraltogether.
9 Whenaconc:entratedsolutionofsucroseisseparatedfromadilutesolutionofrncrosebya
partially permeable membrane, whi(h solution:
a hasahigherconcentrationofwatermolecules
b haslowerwaterpotential
c willexperienceanetgainofwatermolec:ules:1
Thilexperimeotcanbe
da,sorurnoo
b~gcootaining sucrose
sollflooni,Jowered
intowate,;bagshows
no signs of internal
pressure,i.e .. rtisina
flacc id cood,tion
IdJ
carriedoutinyoorlabofato<y
uptakeofwate,
byo,mosis<We,time
dialysis tubingbagnowsttetchedbyhighintemalp,._..sure
(l,y<lmsta ticpressure)duetowate,uptal<e,i.e.itisjn
, ~ =•»rneo<rn••.
ru,,;d rnod"=
owingtothehydrmtaticpressure
exefledbythedialysisbag,whic:h
cannotbestretchedanymOfe
I
thep, ... surepotential{11<,.}
has offsetthesollllepotential{lJI,)
Figure4.12f'r'-"surepotentialatwork
84
4.2 Movement of substances into and out of cells I
Wat er pot ential = solute potential + pressure potential
The concepi of 'water po!ential' allows the effeas of the two fuaors acting on water in a system
(suchasadialysistubing'ba g'oralivingcell) tobebrou ghttogecherinasingleequation:
.,
water potential - soluce potential + pressure potential
We have seen that the water po!ential of a solution is the name we give to this tendency of water
molecules to emer or leave solutions by osmosis, and that the Greek letter psi (symbol ,;,) is used
torepresentthewaterpo!ential.
Since 'water po!ential' is really a measure of the free kinetic energy of the water molecules, pure
water obviously has the highest water po!ential. By definition, this is set al zero. We have also seen
10 Whenaconcentrated that once a solute is dissolved in water, the water mole,:ules are immediately less likely to diffuse
solution of glucose (they are less mobile ). So. the effect of dissolving a solute in water is to lower its water po!ential.
i15eparatedfrom Consequently solutions at atmospheric pressure have a negative value of water po!ential. For
a dilute solution of
example,asolutioncontaining:
glucose by a partially
permeable membrane, JA2g of sucrose in a l()()cml has a water poiemial of - 270kPa, and
whichmlution }4.2g of sucrose in a l()()cml has a water poiential of---J510kPa .
a hasahigherwater
potential Donegativeval,,e.slibetbesegive>''"'f>roblems?
b hasahigher Not really, perhaps. You use them in daily life. Imagine an international weather forecast that
concentration of
repons the temperature in Siberia has changed from - 10°C to - 25°C. You immediately know this
water molecules
means it's much colder there. In other ,vords, - 25°C is a much lo wer temperature than - 10°C, even
c willshowanet though25isalargernumberthan 10.
gain of water
Jn the same way - 3510 kPa is a smaller water po!ential than - 270 kPa, although 3510 is larger
molecules?
numberthan270
Osmosis in plants
We need to think about the effects of osmosis in an individual plant cell first (Figure 4.13). There
aretwofacrorstoconsider.
• In a plam cell there is a cellulose cell wall . This is not presen! in an animal cell .
• In any cell the net direction of water m01-·ernent depends UJXXl whether the water poiential of
thecellsolutionismoreorlessnegativethanthewaterpotentialoftheexternalsolution
When the =temal solution is lcs~ n cgali\•c (cha! is, there is little or no dissolved solutes), there
is a net flow of water Im o the cell . The cell solution becomes diluted. The volume of the cell is
e xpanded by water uptake. As a result, the cytoplasm may come to press hard against the cell wall .
If this happens the cell is described as 1urgld. The pressure poiential that has developed (due to
the stretching of the wall) offsets the solute poiential of the cell solution completely and further net
uptake of water stops. Incidentally, the cell wall has aaually protected the delicate cell contents
fromdamageduetoosmosis .
When the external solution is m ore n cga tl \'c (that is, much dissolved solutes), there is a net
!low of water o ul of the cell. The cell solution becomes more concentrated. As the volume of cell
solution decreases, the cytoplasm starts to pull away from parts of the cell wall (contaa with the
cell wall is maintained at points where there are cytoplasmic connections between cells). The cells
are said to be p las m o lyscd (/ysis means splitting) and are flaccid.
85
IJ Cell membranes and transport
whentheextemalwaterpotentialisless
negativethanthewaterpotentialolthecell
cell solution
diluted
net inflow
of water
cellwallstretchedandthe
cellissaidtobeturgid
(preventsruptureofthecellsurfacemembrane)
theextemal~utionisles,
sameasthewate,potentialofthecell
theexternalsolutiooisthesame
~o~;; ,
when the external water potential is the when the external wate, potential is mo,e
negativethanthewate,potentialofthecell
net outflow
cytoplasmpullsawiljfmmthecellwall;
thecellissaidtobeplasmolysedandis
flaccid
theexternal~utiooismon,
concentrated than the cel l coocentration asthecellsolutioo concentrated than the cell
~ution(a hypotonic solution) (an isotoni c solution) ~utioo(a hype rtonic solution)
Figure4.13Aplantcell inchangingextemal~uhom
l
fina lly.lengthsofir>dividual
cylindersarere-measu,ed.
andchangeinlengthexpressed
asa%oftheoriginallength
Figure4.14Measuringwaterpotentialofplanttissue
86
4.2 Movement of substances into and out of cells I
Steps to the experiment:
Tab le 4. 2Thewaterpotentialof • Immerse (at least) one cylinder in each of the range of live sucrose solutions, 0.2moldm-J to
sucrosesolutioos l.Omoldm-J in a tube. Leave them immersed for one day. Set up a table to record the lengths of
the tissue cylinders from each of the live tubes.
Sucrose Water potential
(moldm-1) (kPal • Re-measure the length of each cylinder and record this in your table. Calculate the change in
lengthasapercentage oftheoriginallength.
• Plotagraphofthepercentagechangeinlengthagainstthemolarityofthesucrosesolution. Read
offthemolarityofsucrosethatcausesnochange inlengthofthetissue.
• Using the conversion table below, quote the water poiential of the tissue you have investigated.
Inevaluatingthe=perimem,coruiderthesignilirnntpo1eruialcausesoferrororinaccuracyinthe
technique. What ;,,,provemet1/s could you make?
'f'p•600 'f'p•900
The water potential of each cell can be calculated ('f' • \II, + \l'p):
'f'dcellA • ---800kPa
'f'ofcell B • - 1200kPa
So net water flow is from cell A to cell B until an equilibrium in water poientials is established.
When we disa,ss water movement through plants (fopic 7, page 135) we will see the flow of
water from cell ro cell in roots. From roo1 hair cells, where water uptake occurs from a soil solution
(of high water poiential, i.e. Jess negative 'f'), to the cells at the centre of the roo1 (of lower water
poiential.Le.morenegative'I').
87
IJ Cell membranesand transport
In the plasmoly,,ed cell, the wall =ens no pressure at aU. As a plasmolysed cell starts to take up water
thecontentsenlarge.Atthepointwherethecytoplasmstanstopushagainstthe,vall.3slightpressure
potentialisproduced(Figure4.16). Aswa!eruptakecontinues.!hepressurepotentialc,ventually
becomes large enough to reduce the cell "s water potential to zero. At that point, water uptake stops.
r5
'"""""ffi'"' ( )
•solute potential;
A
V
wate,uptake
{dilution of cell solution) fullyturgidcell
{pressure potential
-~'"' "'~"''
··"-""''·''
~:~': :;.: () 0
p,es,urepotential•O)
+
agaimt~cellwa ll t
'pressure potential
iJ1Crea,;ngcellvolume high
Figure4.16Changingwaterpotentialolaplantcell
Figure4.17Wilting
88
4.2 Movement of substances into and out of cells I
Osmosis in animals
An animal cell lacks the protection of a cellulose cell wall. Consequently. compared with a plant
cell. it is very easily damaged by changes in the external solution. If a typical animal cell is placed
in pure water. the cell surface membrane will break apart quite quickly. The cell will have been
destroyedbythepressurepotentialgenerated.
Figure 4.18 shows what happens to an animal cell (a red blood cell) when the =temal solutiOfl
h:isawaterpotentialth:itislxxhlessnegativeandmorenegativethanthecellsolution.
Jn most animals (certainly in mammals) these problems are avoided because the osmotic
concentration of body fluids - the blood plasma and tissue fluid - is very precisely regulated. The
water potential is regulated both inside and outside body cells (to maintain isotonic conditions)
This process is known as osm orq,.,,la1ion (ropic 14).
Figure4.18Aredbloodcellin
c:har,g ingextemalsolutions
w hen th e externa l wate r w he n the externa l water when the extern a l water
pote ntia l is less negative pote nti a li s thesameas the potential is more negative
tha nth ewater potentia l wat e rpote ntia lofth ecell, th a nthewat e rpotenti a lof
of the cell e.9.intfiebloodplasma th e cell
I 420kPa
Willnetwaterflowbe
fromcellMtocellN7
:=:::::.
-~-~~ cellsurlacemembraoe
and~lhene,pelled
fromlhecellbylhe
:~...,...JCtk>nof
Explain your answer. Figure 4.19 The role of the ccntfactile vacuole in Amoeba
89
IJ Cell membranesand transport
Figure4.201-iowpola,w~te,molec:ulescrossthelipidbilaye,
90
4.2 Movement of substances into and out of cells I
3 Ac tive trnnspon in,·olves s pecial m o lecules o f the membrane called 'pumps '
lbe pump molecule picks up particular molecules and tr:msporu them to the other side of the
membrane where they are then released. The pump molecules are globular proieins that traverse
the lipid bilayer. Movements by these pump molecules require reaction with ATP. By means of
this reaction, metabolic energy is supplied to the process. Most membrane pumps are specific to
panicular molecules. This is the way selective transport is brought about. If the pump molecule
forapanicularsub.stanceisnO!present,thesubsrnncewillnotbetr:msponed.
Active transport is a feature of most living cells. We meet =amples of active tran.spon in the gut
where absoqxion occurs, in the active uprnke of ions by plant roois, in the kidney tubules where
urine is formed. and in nerves fibres where an impulse is propagated . We discuss some d these
examples later.
molecules
robe
tunsported
into cell
ADPa ndP;,eleased
frnmcarrierprntein,
()==::::::: :::::::,,:::Q which,evertstothe
Q:== :e=o receptMe shape
~
~
o.=
o.=
:;;,,-0
=O
Figure4.21Acbvetransportofasinglembstanc:e
The pro(ein pump, of cell surface membranes are of differeru types. Some transport a particular
molecule or ion in one direction (Figure 4.21). Occasionally, two substances are transported in the
same direction: for example, Na+ and glucose. Others transport two substances Oike Na' and IC') in
opposite directions (Figure 4.22).
91
IJ Cell membranes and transport
A case study: the structure and function of sodium-potassium
pumps in axons
A nerve impul.se is transmitted along the axon of a nerve cell by a momenrnry reversal in electrical
pcxentbl difference in the axon membrane, brought about by rapid movements of sodium and
pcxas.siumions. YoucanseethestructureofanervecellanditsaxoninFigure 15.J,pagejlj.
Sodium-potassium pumps are globular proteins that span the axon membrane. In the
preparation of the axon for the passage of the n=t nerve impulse. there is active transport of
pcxas.sium (K*) ions in across the membrane and sodium (Na+) ions out across the membrane. This
activity of the Na•-K' pump involves transfer of energy from ATP. The outcome is that pcxas.sium
and sodium ions gradually concentrate on opposite sides of the membrane. The steps to the cyclic
actionofthesepump,are·
I With the interior surface of the pump open to the interior of the axon. three sodium ions are
loadedbyatmchingrospecificbindingsites.
l lbe rC3crion of the globular protein with ATP now occurs, resulting in the attachment of a
phosphate group to the pump protein. This triggers the pump protein to close to the interior of
the axon and open to the exterior.
J lbe three sodium ions are now released. and simultanN>Usly. two potassium ions are loaded by
atmchingtospecificbindingsites
4 With the pcxassium ions loaded. the phosphate group detaches. This triggers a reversal of the
shape of the pump protein; it now opens to the interior. again, and the pcxas.sium ions are
14 Sample5offiveplantti1suediskswereincubatedindilutesodiumchloridemlutionat
different temperatures. Aher 24 hours it was found that the uptake of ions from the
solutionswere{inarbitraryunits)
1,.,~a<S°C
~°'~""°'
Comment on how absorption of sodium chloride occurs, giving your reasons
92
4.2 Movement of substances into and out of cells I
Movement by bulk transport
Bulk transport occurs by movemems of \'Cs idcs of maner (solids or liquids) acros, the cell surface
membrane. The flexibility of the fluid mosaic membrane makes this activity possible, and energy
from metabolism (ATP) is required. Figure 4.23 .shows this form of transport across a cell surface
me b, ~e, ·n arion.
15 Phagocyticcell1are
also found in the
airwaysofthelun!J:5
_s_-" ,
eo<=<~ c ~ O o q , ~ " ' ~ ' " " " " ' ''""'""
Figure4.23 MoYementbybulktransport
Vv'hat istheirroleand Exoq101ds i., !he process by which cells =port products such as enzymes by means of vesicles. We
howdotheycarryit have seen that vesicles maybe budded off frcm the Golgl apparatus (Figure 1.16. page 18). lbe
="
16 Oistinguishbet11<'een
vesicles are !hen guided to the cell surface membrane by !he network of mkrorubule-s in the cytoool.
Here tOC,y merge with the membrane and their contents are discharged ro the outside of the cell.
thefollov.ingpairs . Alternatively, wasce maner may be disposed of at the cell surface, in a similar way.
a protein1andlipids By the reverse process, cndocyto,;L'<, substances may be imported into the cell. Here, fresh
in cell membranes vesicles are formed at the cell surface. This occurs when part of the cell surface membrane forms
b ac:tivetransportand a 'cup" around a particle or a drop of fluid. and encloses it to form a vesicle. This vesicle i'l then
bulk transport broughtintothecytosol.
c hypotonicand The whole sale import of solid maner in this way is termed phagocytos is. In the human lxxly,
hypertonicsolutions there is a huge fon::e of phagocytic cdls , called the macrophages. Some of these mop up debris of
d eodocytosisand damaged or dying cells and break it down . For example, we dispose of about 3 X 10" red blood
exocyto1i1 cells each day! All of these are ingesced and disposed of by macrophages.
l'inoc}1t"' L~ is bulk import of fluids and occurs in the same way. Bulk uptake of lipids by cells
lining the gut occurs as part of the digestion process. for example.
Exocytosis:secretionofmaterialsoutofcellsbycytoplasmicve1iclesfusingwiththecell1urfac:e
membrane and releasing the contents of the vesicle into the fluid around the cell, using ATP to
movethecytoplami
Endocytos is: uptake of materials into cells by inward folding1 of the cell MirfilCe membrane to
form sacs of membrane that separate from the membrane to form vesicles within the cytoplasm,
u1ing energy Imm ATP to move the cytoplasm around. The process may involve liquid solutions
or1mpension1(pinocytosis}or1ol idmac:romolecule1orcells(phagocytosis}
93
IJ Cell membranes and transport
:::;;:"'.
,,!'"':'::h,h,o~IO •
allowsooespecifK
substance to pass
0 0
o •-+I _. •.•
~
~,:',:":~:ne -
energyfromATP
•
• '---"
+
isusedselec:tivelyto
mo~one(ortwo)
spec:ifKsuhstar,ces
+ ____.,~
ATP
ADP+Pi
suhsttatemolec:u~
fit<hereandthe
,eactKlf\thenoccu"
cell-<ellrecognitionsite -
attachmentmay,esultin
cells binding together
binding sites {
forantigen-antibody,eactKlf\
(Topic:11)
94
Summa~ •
• The cell surfa(e membtane isan organellethat~nds • The name given to the tendency of water molecules to mOYe
and containsthece ll cOl'lle nts. Aoossilthereisacontinuous about is water potential. TheGreeklelterpsi(symbol.-)
move ment o f t he mole<ules needed by cell§ a being is used to represent water potenti<ll. Water moves from a
di~ofbythem. regionofhigherwaterpotentialtoaregionoflowerwater
• The membrane consists of lipids a nd prote ins with a small potential- do.vn a water potential gra die nt. Equilibrium
amount of carbohydrat e . The lipids are phospholipids, is reached only if or when the water potential is the same in
~eswithahydroph~icheadandahydrophobictail, both regions. At this point. there would be no net movemeot
Dfganisedintoilbilayerwiththeheadsontheoutsides.The of water from one region to another. but random mcwemenl5
proteinsare globularprote ln sarrangedin,acrossandon of water molecules continue
thesurfaceofthelipidbilayer. Shortcarbohydra te chainsare • Solutemolecules insolvtionaresurroundedbywater
attachedtosomeoftheproteinsand(pidsontheoutside molerulesheldby hydrogenbond s.lnaconcentrated
lnaddition,avariableqoantityofmoleculesolcholesterol solution,fewwatermolecules¥efreetomove. Thereforethere
occurs,insertedbetweensomeofthelipidmolecules. isnetdiffu sionof freewatermolecul esinpurewaterora
• The membrane is described as a fluid mosaic, as the dilutesolution(freewatermoleculesinhighooncentration),
components(particularlythelipids)moveaboutlaterally, throughapartlal lypermeablemembrane (suchasthecell
relatiVi:'toea,chother.andtheproteinsappeartoberandomly rurfacemembrane),intoamoreconcentrated solution{where
scattered about. Thecholesterolregulatesmembranefluidity. freewatermoleculesareinlowconcentration}-aprocess
kno.vn asosmosls.
• There an:- three mechanisms o f transport across
membra~. In diffu sion (and osmosi s, a speciill ca5e of • Activeuptake isaselectlveprocessandinvotvesthe
diffusion) the energy comes from the kinetic e nergy of matter. accumulationofionsandorganicsubstancesa galnst
In active transport (l .e. pumping) and in bulk transport, the a concentration gradient. The process is powered by
energy is pl'Ollided ifldirectty from cell metabolism, via ATP. metabolicenergyandinvolveshighlyspecific prote in
molecules in th e m e mbran e that act as pumps.
• Diffusion occurs from a1egionofhighconcentrat ion
toaregionoflowooncentratiof'I. The lipidbilayerofthe • Bu\ktransportisthe1T10Yementoftheoontenl5ofvesic.les
cell surface membrane acts as a barrier, a t lei151 lo polar (t iny,mefTID"ane-boundsacs)ofmatter,solidor liquid,into
substanc:es. Polarsubstancessuchaswaterdiffusethrough (e ndocytosis)oroutof{ekocytosb)thecel
tiny pores or small spaces in the bilayer. Some molKU!es
reactwithamembraneproteintoopenaspecificchannel
(facilitated diffusion).
95
IJ Cell membranes and transport
4 Designandoutlinealaboratoryexperimenttoinvestigatethe
effect of temperature on the cell rnrface membrane
Usesample1ofplanttissuethatcontainsanintensely
red,water..,.;olublepigmentinthecellvac:uoles{suchas
beetroot}
a)What1tepswouldyoutaketoen!.llrethesignificanceof
Fig. 2 .1 yourresults7
b) What outcomes to your experiment do you expect7
a)WithreferencetoFig.2.1 and the information above,
explain h<M' flatworms survive without a transport system
for respiratory gases [4]
96
AS Level
97
II The mitotic cell cycle
generation to generation during reproduction. Both of the functions depend on "informatico'. The
information in the nucleus is contained within structures called ch romosom es. These uniquely:
e arecopiedfromcdltocellwhencellsdivide
• arepassedintonewindividualswhensexcellsfusetogetherinsexualreproduction.
So. the nucleus ccotains the chromosomes of the cell. and the chromosomes contain the coded
instructicos for the organisation and activities of cells and for the whole organism. It is on the
structure of the chromosomes that we focus first.
The information the nucleus holds o n its chromosomes exists as a nucleic acid called
deoxyribon ucleic acid (DNA). DNA is a huge m olecule made up of two paired strands in the
form of a double helix (Figure 6.J, page l 13). A single, enormously long DNA molecule runs the
full length of each chromosome.
Lookuptbestruc/ureofDNAnow
Too c l , - ~ a.., effectively a linear series of genes. A pan:icular gene always occurs on the same
c l , - - in the same position. We can define 'gene" in different ways. For example, a gene is·
• a specific region of a chromosome that is capable of determining the development of a specific
characteristic of an organism
• a specific length of the DNA double helix, hundreds or (more typically) thousands of base pairs
long,whichcoclesforaprotein
e aunitofinheritance.
Meanwhile, there are four features of the chromosomes that are hdpful to note at the outset:
• "Jhe s hape of a d u o m o,;.om c is c h arnctcristk . O,romosomes are long, thin structures of a
pankular, fixed length. Somewhe.., along the length of the chromosome occurs a shon, constricted
region called the cen trom crc. A centromere may occur anywhere al cog the chromosome, but it is
always in the same posi!ico on any given chromosome. The position of the centromere, as well as
the length of a chromosome is how they a.., identified in plxxomicrographs
• The nrunbcr o f c h ron i.-.m cs per species ls fi xt.-d. The number of chromosomes in the
cells of different species varies. but in any one species the number of chromosomes per cell
is n ormally constant. For example, the mouse has 40 chromosomes per cell. the onion has 16,
humans have 46, and the sunflower J4. These are the chromosome numbers for the species.
• d 1rom osom es occur in p,-tlrs . The chromosomes of a cell occur in pairs, called homologous
pairs (meaning 'similar in structure"). One of each pair came originally from each parent. So, for
example. the human has 46 chromosomes, 23 coming originally from each parent in the process
of sexual reproduction. This is why chromosomes occu r in homologous pairs
• d 1rom osom es are cop ied. Between nudear divisions. whilst the chromosomes are uncoiled
and cannot be seen. each chromosome is copied. This copying process occurs in the cell
before nuclear division occurs. The two identical structures formed are called chrom a tlds. The
chromatids remain attached by their centromeres until they are divided during nuclear division.
TI>en the cc nt rom crcs and the chromatids separate. After this separatico, the chromatids are
recognised as chromosomes again. Of course, when chromosomes copy themselves the critical
event is the copying of the DNA double helix that runs the length of the chromosome. This is
known as replication of the DNA. You will discover how this replication process i, brought
about in Topic 6.
98
5.1 Replication and division of nuclei and cells I
The cha nging a ppeara nc eo f a chromosome
inthe nu cle us, not after th e chromosome has asaconsequ e nceofn uc lear
yet duplicat ed madeacopyofitself divisio n(mitosis)
Well, take the case of human DNA . In each nucleus, Ule torn] length of the DNA o f Ule
d r r - = is over 2 metres. We know Ulis is shared out between 46 chromosomes.
O,romosomes are of different lengths, but we can estimate that within a typical chromosome of
5µm length, there is 3 DNA molecule approxillllltely Sctn long. This means that 3bout SOOOOµm o f
DNA is packed into 5pm of chromosome.
This phencmenal packaging problem is 3chieved by the much-eoiled DNA double helix being
looped 3round proi:ein beads o:insisting of a packaging protein rnlled histon e . This is 3 basic
(positively crulrged) proi:ein conmining a high COflcentration of 3mino acid residues wiUl additional base
groups (-NH,), such as lysine and arginine. Eight histooe molecules combine to make 3 single be3d.
Around each bead, the acidic (negatively charged) DNA double helix is wrapped in a double loop.
At times of nucle,ir division the whole be3ded thr"3d is itself =iled up, fonning the chromatin fibre.
lhe chro!lllltin fibre is again coiled, and the coils are ]coped 3round a 'scaffold" protein fibre, made
of 3 non. llis tone pro ld n . This whole strum1re is folded (supercoiled) into Ule much-condensed
c h = ~ e , shown in Figure 5.2. This arrangement eruibles the 53fe storage of these phenomenal
lengths of DNA trult 3re packed in the nudei. However, it also allows access to selected lengths of the
DNA (particul:ir genes) during transcription - 3 proces..s you will encounter in Topic 6.
99
II The mitotic cell cycle
Hffl!tM1truchlrt is~unpa,d<Nl
tosllOWl'OWilhugemolKu~ofDNAi,r.eld
andsupported,by.
1)double-cO<lngarO<ndhistooeproteim
(be~WucMts•nudtosome,)andthen
2)1oopedalong!Mleogthof thechromat,d,
• l• ctronmkrogrilphof<hrornosom• attached1oaproteinscalfokl
durlngmltosis{metapha,e) -,howiog
twodvomatldsheld log<'lhl>fat the
centrome<e(~400Xl)
(typblyaONAmolKule
olaboutSaninlfngthis
p,ckedintoNChchrornatidol
;,ppro>Omiltely5JLITlinlength)
AlsopresentilfE'eraymes
inYolvedin!hetran'lCriphon
and replication d tM DNA
<Xlftol nud ..,"""'•
olei!1,t his!one
~~
Figu,.. 5.2 The pK~1119 of DNA in the chromosome -the role ol t.,tone protein
100
5.1 Replication and division of nuclei and cells I
This is the arrangement in the nuclei of eukaryOles. We can conclude th3! the single circular
d r r = = ofprokaryoies does not present the same p ackaging problem. for in these org3nisms
2 Deducethesignificance theDNAisdeseribedas ' naked'. ltoccurswithoutthesupportofanyprotein.
ofthepo5itivelycharged
histoneproteinandthe The nature and significance of telomeres
negatively charged ONA
Each time the DNA of the chromosomes is copied (replicated), prior to mitosis and cell division,
the ends of the DNA molecule s cannOI: be copied. As a result, a few terminal nucleotide sequences
are lost. With repe-:,ted replications o:mes progressive shonening on the DNA of e:ich chromosome.
If the ends were nOI protected in some way. chromosomes would Jose genes or parts of genes. Jn
facttheendsareprotectedbyspecbl.non-codingnudeotidesequencescalledtelomeres. Typically,
a telOfllere consists of 3 six-nucleotide sequence such a., TIAGGG" , repeated up to a thousand
times. h is parts of this non-coding sequence that 3re lost at each replication of the DNA, without
anyh3rmtothegenesthechromosomecarries
Jn mltoisis . the daughter cells produced have the same number of chromosomes as the parent
cell . typically two of each type and this is known as the diplo id (2 ,.) state. Mitosis is the nuclear
division thm occurs when cells grow or when cells need to be repbced and when an organism
reproducesasexu3lly.
Jn mcloisls, the daughter cells contain half the number of chromosomes of the parent cell . Th3!
is. one chromosome of e:ich type is present in the nuclei fanned and this is known as the
haplo id ( ,.) state. Meiosis is the nuclear division that occurs when sexual reproduction occurs,
rypically during the formation of the sex cells (called gam e tes) (see pages 345---8).
Whichever division takes pbce, it is normally followed by division of the cytoplasm to form
separate cells. Thisprocessiscalledcytoklncsis.
101
B The mitotic cell cycle
3 a Whati1ahaploid
cell/
b Whereinthehuman
body would you find
Fo\lfcellsformed,eachcootainingonly
cells undergoing onesetofch-= (onlyoneof
i mei05is eac:hhomologeousch,==) -
ii mit05ois7 haploid (nl
Figure5.4Mitmisandmeiosil:thelignificantdifferl'J1Ces
lnterphase
lnterphase is always the longest pan of the cell cycle but it is of extremely \'ariable length . At
first glance the nucleus appears to be 'resting' but this is noc the case at all. The chromosomes,
previously visible as thread-like structures, have become dispersed. Now they are acrively involved
102
5.1 Replication and division of nuclei and cells I
the cell cycle ,,, change in ce ll volumeand quantityo fDNAduring ace ll cyc le
mitosis and
cell division
FigureS.SThestagesofthecellcyde
in protein synthesis at least for most of interphase. From the chromosomes, copies of the
information of particular genes or groups of genes are taken in the form of ribonucleic acid (RNA),
('messenger RNA", page 123), for use in the cytoplasm. In the cytoplasm. ribosomes assemble
proteins from amino acids, combining them in sequences dictated by the information from the
gene.Wererumrothisproces.slaterinthistopic.
Look at Fif!ure 55. lbe sequence of the events of interphase is rnustmted there. lbe changes
thatoccuraresummarisedinTable5.I.
Table5.1Theeventsolinterphase
103
II The mitotic cell cycle
Cancer - diseases of uncontrolled cell division
CanceristheresultoftheUJ-regulatedproliferationofcells,typicallyresultingintheformation
of a solid tumour. There are many different forms of cancer. affecting different tissue., of the
body - cancer is not thought of as a single disease. Cancers are usually fatal if untreated . Today,
many cancers are treatable , and in these cases. survival rates are improving markedly.
Today, in developed countries, Ofle in three people suffer from cancer at some point in their life
and approximately Ofle in four will d ie from it. ln these regions the commonest cancers are of the
lung in males and of the breast in females . However. in many paru of the world. cancer rates are
different - often they are significantly lower. You can Moe the range of common cancers and their
incidences worldwide at: lutp,//globocan.iarc.fr/
So,cancerarise-swhenthecellcycleoperate-swithoucilsnormalcontrols. lnahealthyceU
the cell cycle is regulated by a molecular cOfltrol system, itself controlled by specific genes. However,
cells may start ro divide by mitosi'l , repeatedly, without control or regulation. Now the rate of cell
multiplication i'l very much fas!er than the rate of cell death, arx:l an irregular mass of cells, a rumour,
i'lfrnrned(Figure5.6)
~::.§3--=::~~~
to the DNA molecules of chromosomes. The result is a
mutation - achangeintheamountorchemicalstructure
of DNA of a chromosome. Mutations of different types
build up in the DNA of body cells over time. A single
mutationisunlikelyroberespon.siblefortriggering
abnom,alcells
(donotcauseserious
rapid
m,tosis
I cancer. With time, mutations may accumulate in the body
and then trigger a disease. This =plains why the majority
of cancers arise in older people. Also, mutation may
occur in the cells of different body tissues, and hence
::..mb.!::i"t,c:,ge,y~ • f
-\(.e; cancer·snotones"ngled"sease.
fl .~,:. Whateverthecause,twotypesofgenesplayapartin
!:ye~~!~!:~~~:= ;;_u,'.;,~~;nlarges, '•••~~ initiatingacancer,iftheymutate:
doesnotspreadfromotssiteof ,rntiaboo . • Proto-on cogen es. These are genes that code for
theproteinsthatplayapaninthecOfltrolofthecell
cycle. When a prot0-0ncogene mutates. an oncogene
:~igh':v~t~~=; cells ;~:,is
results. lnthisstate,thegenepermanentlyswitches
Thisstepispossible '
butnot;nevitable ,
Amalignanttumour consi<;tsolcellsthat
,ec,etesignalstngge,;r,ggrowthofbkioda<>d
in DNA when they arise-, or instruct the cell to die
(by programmed cell death). These genes. too, are
vulnerable to carcinogens. For example, a mutation
.•• ~hvesselstose,vethetumoo, cellsa tthe
' ·~=•of""'""=· A~chmffio<o mayinactivatethegenethatcodesforapro1einknown
~~; s:.·::,:.'::~:::: ::::,b:::;;;;;:'.,':!' as p53. When protein p53 i'l present, copying of faulty
DNA is stopped. Then, other enzymes are able ro
:~::.% i~~i:;:~~:,:~~ed repair the DNA and correct the fault, or programmed
celldeathi'lswitchedOflinthefaultycell.socancer
,eadofcaocercellstootherlocations is avoided. Abnormalities in the p53 gene have been
found in more than 50 per cent of human cancers.
Figure 5.65tep,;nthedevek,pmentofamalignantlumoor
104
5.2 Chromosome behaviour in mitosis I
Table S.2 Common can: ir,ogem known to increase mutaboo rates and the l,kl'lihood of carn:er
In m eta phase the two centrosomes move to opposite ends of the cell . Microtubules of the
cytoplasm stan to fom, into a spindle, radiating out from the centrioles (page 19). Each pair of
chromatids is attached to a microtubule of the spindle and is arranged at the equator of the spindle.
(Note: in plant cells, a spindle of exactly the same structure is formed but without the presence of
thecentroscmes).
6 A5thecontent1ofa Jn a n a ph asc the centromeres separate , the spindle fibres shonen and the chromatids are
nucleus in the human pulled by their centromeres to opposite poles. Once separated, the chromatids are referred to as
body prepares to
undergom itosis,how
Jn 1doph asc a nuclear envelope reforms around both groups of chromosomes at opposite ends
manychromatidsdoes
it contain? of the cell . The chromosomes 'decOfldense" by uncoiling, beo:>ming chrom:,tin again. lbe nucleolus
reforms in each nucleus. lnterphase follows division of the cytoplasm.
105
II The mitotic cell cycle
Cytokinesis
Cycok.inesis is the d ivision of the cytoplasm that follows telophase. During cytokinesis, cell
org:rnelles such as mitochondria and chloroplasts become evenly distributed between the cells. In
animal cells, the cytoplasm separates by in-tucking of the cell surface membrane at the equator of
thespindle,'pinching'thecytoplasminhalf(Figure5.8).
Jn plant cells. the Golgi apparatus forms vesicles of new cdl wall materials which collect along
the line of the equator of the spindle, known as the cell plate. Here the vesicles merge, fonning the
new cell surface membranes and the cell walls between the two cells (Figure 5.7).
ERsecretesYe<iclesof
wall fom,mgenzyme,
remains of the - - + ! - ,~ ~
sp ind~
Golgi"!)paratus
secn,tesvesicles
c::::;> daughter cells
of wall compor,ent,;/"tif-"r\:'",-',c-i._;_J-"
cellp!ate
(equator of ~rnllectat themidlrne
spindle)
thenfusetofrnmcellwall
laye" and new cell surface membrane
Figure 5.7Cyto+:inesisinaplantcell
Mitosis - a summary
Daughter cells produced by mitosis have a set of chromosomes identical to each other and to the
parent cell from which they were formed because:
• an exact copy of each chromosome is made by accurate copying during interphase, when two
• chromatid-. remain a u ached by their centromeres during metaphase of mitosis, when each
becomes attached to a spindle fibre at the equator of the spindle
• centromeres then separate during anaphase and the chrornatids of each pair are pulled apan to
oppo!!iile poles of the spindle. Thus, one copy of each chromosome moves to the poles of the
spindle
• the chromosomes at the poles fonu the new nuclei - two to a cdl at this point
• two cells are then formal by separation of the cytoplasm at the mid-point of the cell, each wi1h
an e xact copy of the original nucle u s.
7 IJ:<.ingslOOtheyhadpreparedtoobservechromosomesduringmit~inaplantroottip(Figu!l'
-..
S.9),fivestudentsobservedandrecordedthenumberofnudeiateachstageinmitosisin100cells
Number of nuclei counted
~~:' 1Student2 I Student) 1 Student4 1 Student5
I" l ,s I" In
Meta phase Im 1, In 1,
Ana phase 1, 1, 1, 1,
Telophase I» l >e In In
a Calculate the mean percentage of dividing cells at eilCh stage of mitosis. Present your
resultsasapiechart.
b Assuming that mitosis takes about 60 minutes to complete in this s.pecies of plant, deduce
whatthe~resultsimplyaboutthelengthsofthefoursteps?
106
5.2 Chromosome behav1ourin
. mitosis I
equatorial
plate
~ -
-1 chmmat>ds
pulledapa<t
bymocrotubute<
u!J ~-,anat,dsj(>ned
v1Cl'll!mmere
:;:,:::e~:i~
Figu,..,5.8Mitosisinananimalcell
107
II The mitotic cell cycle
Figure5.9Prepam,ganonionroottipsq"'1shwithethano-orce,nstain
108
Examination style questions I
• The nu cle uscontrolstheilclivitiesofthecellthroughoutlife lengthsoftheDNAcodeforparticularpolypeptidesandare
ltisthelocationofthehereditarymaterialwhichispassed called ge nes
fromgenerationtogenerationinreproduction. The division of • Re pli ca ti o n o f the DNA of the chromowme1 occurs in the
thenucleusisthefirststepincelldi visio n in terph ase of the cell cycl e when the chromosomes are less
• Chro mosomesoccurinthenudeusbutareonlyvisible, coiled,wellbeforenudeardivi1ionoccurs
when1tained,attimesofnudeardivision.Thenumber • Each time the DNA of the chromo50me1 is copied (replicated}
ofc:hromosomespernudeusineachspeciesisfoced priortomito1i5andcelldivision,apartoftheendoftheDNA
Chromosomesoccurinpairsandhaveacharac:teristicshape. molecules of chromosomes is lost . Ultimately, chromosomes
• M itos is is the replicative nuclear division in which, after cell wouldlosegenesorpartsofgenesbythisproces1,ifthey
division,thenudeuspresentineac:hoftwodaughtercells werenotprotectedbythepresenceoftelo me res. Telomeres
formed has exac:tly the !idme number of chromosomes as the arelong,seeminglydisposablenon-codingsequencesof
parentcell, typicallytwoofeac:htype. ltis in the di plo id (2n) nucleotidebasesattheendsoftheDNAmoleculesof
state. Mitosis is am>eiated with cell divisions of growth and chromosomes that fulfil this important function
asexual reproduction • Mutatio nsareunpredic:tablechangesinthegeneticmake-
• Nucl e ic aci ds are long, thread-like macromo lecul es with a upofacellinvolvingthesequenceofparticularbasesatone
uniformandunvarying'bac:kbone'ofaltematingsugarand loc.ation(genemutation)
phosphatemolecules.Attac:hedtoeachsugarmoleculei1a e Ca ncersareinitiatedwhencellsstarttodividerepeatedly,
nitrogenous base 1\t.ich projects sideways. Since the bases by mitosis, without control or regulation . An irregular mass
vary,theyrepresentauniquesequencethatcarriesthetod ed ofcellsislormed,calleda tumo ur.Therearemanyform1
in fo rma ti o n heldbythenudeic:acid. ofthedisea1e,affectingdifferentti11uesofthebody.Often
• DN A exists as two strands arranged a1 a do ubl e he li x. It cancersaretheresultofnumerouschancemutatiomtriggered
occurs in the chromosomes in a highly coiled state, Mipported by agents known as carcinogens. These include ionising
by a pro t e in fra mewo rk. Within a chromo50me, specific radiation,tobacco1moke
109
AS Level
Und=tanding the structure By the end of this section you should be able to:
ofnudeicacidsall01\!San
understandingoftheirrole a) decribethestructureof nucleotides,i ncludingthephosphorylat«lnud eotide ATP
in the storage of genetic b) decribe the structure of RNA and DNA and explain the importance of bue pairing and the
information and how that differenthydrogenbondingbetweenbases
information is used in the c) describe the semi-con,ervat ive replication of DNA during interphue
synthesis of proteins
110
6.1 Structure and replication of DNA I
The nitrogenous base-s are derived from one- of two parent compounds, purine or pyrimidine. The
purine bases are cytoe;.inc or guanine; the pyrimkline bases are ade nine , thymine or urac il. lbese
molecules are illustrated in Figure 6.1. You do no( ne,,d to know their structural formulae. However,
you should nole that purines have a double ring strucrure and pyrimidines have a single ring
structure. Wewillno{ethespecial significanceofthissizedifferenre,shortly. lthasbecometraditional
in biology ro abbreviate the names of these five bases ro their first leners, C, G, A , T and U.
The three components, nitrogenous base, pentose sugar and phosphoric add, are combined
by condensation r eac tions to form a nucleotide with the formation of two molecules of
water. The structure of these components and how they are combined together are illustrated
in Figure 6 .1. Here a di3grammatic way of represenrntion has been used to emphasise their
spatial arrangement.
the components:
phosphoficadd
nitrogenous bases
I"""~ l
0~'!5
phosphoticac:,d shownd iag,ammaticallyas
o~~
r o, "i,, J-, *
f8 phosphate
suga, (pentose)
N:..J-NHi
1 Oistinguishbetween
a'nitrogenousba5e'
base ob...
OH OH
anda 'base'foundin
~-
ino_,,..
_ ,_• _me
_ m_ist_o _. ~ fig i.-e 6.1 The compor,ent,; ofmKlootides
111
II Nucleic acids and protein synthesis
of chemical energy locked up in its struaure. What makes ATP special as a resen,oir of st<>ttd
chemical energy is its role as 3 common intennediate between energy-yielding reaaions and
energy-requiringreaaiOflsandprocesses.
Energy-yielding reaaions include the photophosphorylation reactions of photosynthesis
(page '11:,7). and many of the reaakos of cdl respiratiOfl (page 243).
Energy..-equiringreactionsincludethesynthesisofcellulosefromglucose,thesynthesisof
proteins from amino acids and the contraaion of muscle fibres.
The free energy available in ATP is approximately 30-J4kJmc..i-', made available in the presence
of a specific enzyme, referred to as ATPase. Some of this energy is lost as heat in a reaaiOfl, but
much free energy is m ade availa ble to do useful work, more than sufficient to drive a typical
energy-requiringreaaiOflofmetabolism.
Direction in the nucl e ic • Sometimes All' reacts with water (a hydrolysis reaaion) and is converted to ADP and Pi. Direa
a cid molec ule : hydrolysis of the terminal phosphate groups like this happens in muscle contraaion, for example.
inanudeicacidmok>cule, • Mostly, All' reaas with other metabolites and forms phasphorylared intermediates, making them
thephosophategroopsare
more reaaive in the process. 1he ph06phate groups are released later, so boih ADP and Pi become
bndgesbetweenu rbon-3of
ooesuga,mok>culeand availableforreusea.smetabolismcontinues.
ca rbon -5of theneJ<t
Consequently,wecanidentify
'direction 'inthenucleicacid Nucleotides become nucleic acid
molec:ule,indicatingoneend Nucleotides themselves then combine together, one nucleotide at a time, to form huge molecules
as5'andtheothe,endas3'
called nucleic acids or polynuc kotides (Figure 6.2). This occurs by a o:indensation reaction
between the phosphate group of one nudeo{ide and the sugar group of ano1:her nucleotide.
Up to 5 million nucleotides, cOfldensed together, form a polynucleotide. So, nucleic acids are
very long, thread-like macromolecules with alternating sugar and phosphate molecules forming
the 'l:xickbone'. This pan of the nucleic acid molecule is uniform and unvarying. However.
also anached to each of the sug3r molecules along the strand is one of the bases, and these
project sideways. Since the OOses vary, they represent 3 unique sequence that carries the coded
informationheldbythenucleicacid.
Introducing DNA
The DNA m o lecule consists of two polynucleotide strands, paired together. 1bese strands are held
by hydrogen bonds between their paired bas<,s. The two strands are of the order of several million
nucleotides in length and they take the shape of a double h e lix (Figure 6.J). In all DNA molecules
thenudeotidescontaindeoxyribose
The Olher chemical feature that distinguishes DNA from RNA is the organ ic bas<,s present and
the way they pair up. In DNA the nitrogenous OOses are C)1oslne (C), guanine (G), adenine (A)
and thymine (0, but never uracil (U). 1be pairing of bas<,s is between adenine and thymine ( A
and "I) and between cytosine and guanine (C and G). This is because there is just enough space
between the two sugar- phosphate backbones for one purine and one pyrimidine molecule. So a
purine in one strand must be opposite a pyrimidine in the O!her. and vice versa. These are the only
combinations that fit together along the helix. Pairing of OOses is accompanied by the fonnatiOfl
of hydrogen bonds: two are formed between adenine and thymine and three are formed between
cytosine and guanine. This pairing, known as complcme n L'lry b a..,;.c pairing, also makes possible
the very precise way that DNA is copied in a process railed replication.
We have noted there is direction in a nucleic acid molecule. Look agai" at Figure 6.2 to make
Whenthepolynudeotide cerlai" y,m ,mderstand what 'direction· means
chaing rowsbyadditionof So, in the DNA molecule, the phosphate groups along each strand are bridges between rarbon-3
anothefnudeobde,the of one sugar molecule and carbon-5 of the next . Funhermore, within the DNA double helix the two
nucleotidesa ,ealwa)'Sadded chainsarearrangedantipa rallcl. JnFigure6.Jyoucansecthatonechainrunsfrom5'toJ'whilst
tothe3'end
the Olher runs frcm 3' to 5' - which is what 'antiparallel" means.
Figure6.2 "0irection'inthe The existence of direction in DNA strands becc.nes important in replication and also when the
nudeicaddmolecule genetic code is transcribed into messenger RNA.
Ill
6.1 Structure and replication of DNA I
inthechromosomes,thehelKal,tructu,eofONA
isstabilisedandsuppottedbypmteim
theDNAmoleculeistwisted
into a double helix
Figure 6 .3The,tnKtU<eOf0NA
Introducing RNA
sugar--p~backbone The RNA m o lec ule is relatively shon in length. compared with DNA. In
fact, RNAs tend to be between one hundred and one thousand nucleotides
I Jong, depending on the panirular role they have. In all RNA molecules the
nucleotides containribose
The other chemical fe'llture that distinguishes RNA is that the bases are
cytOlline (C), guanine (G), adenine (A) and u racil (U), but never thymine
(I). RNAoccursasasinglestrand
Jn the 'information business' of cells there are two functional types of RNA.
l1>ese are rnc.,s;;e n i,.,cr RNA (mRNA) and 1r:111sfer RNA ( tRNA). Their roles
are in the transfer d information from the nucleus to the cytoplasm and in the
sing le strand of cons1ruction of proi:eins at the ribosomes, located in the cytoplasm. We will
polyoudeotkle rerumtothisprocess,shortly.
withribmes.uga,
and nitrogenous
bases:aden ine,
ma 'l,guan ·ne
and cytosine Le ngth Verjlongstrands,several million Rela t,Velyshortstrancls
nudeotideslong
Sugar 0eoxyr;bose
C,G,AandT(notU) C,G,AandU(notT)
Consistsoftwo p0¥'uc:leotidestrands Consistsofsinglestrandsand
inthe/Ofll\o/adooblehelixwith existsintwo functional lonns
complementartbasepai,s • messenge, RNA(mRNA)
• CwithGand A withT • t,amlerRNA(IRNA)
Figure6. 5Thestructu,eofRNA
• heldbyhydrogenbonds
113
II Nucleic acids and protein synthesis
The existence o f the DNA d o uble h dix was discovered, and the way DNA holds information
was suggested by Francis Crick and James Watson in 1953, for which a Nobel Prize was awarded
(Figure6.4),
FrancL-. Cric k (1916--2004) and J ames Watson (1928,---) laid the foundations of a new branch
of biology - cell biology - and achieved this while still young men, Within two years of their
meeting in the Cavendish laboratory, Cambridge (1951), Crick and Watson had achieved their
understanding of the nature of the gene in chemical tenns,
Crick and Watson brought together the experimental results of many other workers, and frcrn
this evidence they deduced the likely structure of the DNA molecule,
• Erwin d 1crgaff measured the exact amount of the four organic bases in samples d DNA,
and found the ratio of A:T and of C:G was always close to 1. D,ergaff's results suggest
consistent base pairing in DNA from different organisms
RatioofbasesinDNAsamples
Organism Adend in e:Thymine I Gua nin e:Cytos.ine
1 1.00
1 1.00
1 1.02
10.99
• Roisalind Franklin and Maurice Wilkins produced X-ray diffraction patterns by bcrnOOrding
crystalline DNA with X-rays.
Crkk and Watson concluded that DNA is a double hdix consisting of:
• two polynucleotide strands with nitrogenous bases stacked on the inside of the helix Oike
rungs on a twisted ladder)
• p.trallel strands hdd together by hydrogen bonds between the p.tired OOses (A- T, C-G)
• tenperrumofthehdix
• twoantiparallelstrandsdadoublehelix.
They built a model. See Figure 6.3 for a simplified model of the DNA double helix.
a) Watson and Cod \\ilh thei, demonstfation model b) Rosalind Franklin produced the cl X-ray diffract>On pattem of ONA
of ONA keyX-,aydiffrac:tionpattemofDNA
atKir,gsCo11ege,London
Figure6 .4Thediscove,yofthero.-ofONA
114
6.1 Structure and replication of DNA I
Replication - how DNA copies itself
Some features of the DNA of the chromosome and how it is copied or replicated have already
• Replication must be an =cremely accura!e process since DNA carries the genetic message.
• Replication is quire separate from cell division, for replication of DNA takes place in the nucleus
during inter-phase , well before the events of nuclear division.
• Strands of the DNA double helix are built up individually from free nuc koiidcs. The structure of
nudeotidesisshowninFigure6.t.
Before nucleotides can be replicated, the DNA double helix has ro unwind and the hydrogen bond,
haldingthesirand.stogethermuscbebroken, alkmingthetwoscrandsolthehelixtoseparate. lhe
enzyme hdkasc brings about the unwinding process and hold, the sirands apan for replication to
occur. Each point along the DNA molecule where this occurs is called a rcplk:uion fork (Figu,., 6.6).
At a replication fork, both strands act as templates. A molecule of the enzyme DNA polymerase
binds to e:ich of the DNA strands and begins to move along them. As e:ich base along a strand is
reached, a free nucleotide th.u is complementary is paired with it (A with T, C with G). lbese free
nucleotides are already 'actiV3ted" by the presence of extra phosphates. Each free nucleotide is held
in place by the hydrogen bonds that form between complementary bases. Then, from each free
nucleotide , the extra phosphates are broken off. This provides the energy by which the remaining
phosphate is joined to the sugar molecule of the neighbouring nucleotide. This condensation
reaction is brougtu about by DNA polymerase. The new strands of DNA 'grow' in this way.
DNA polymerase moves along the single strands of DNA in the S' -4 3' direction . If you look
at the illustration d replication in Figure 6.6, you will see that only one strand can be copied
continuously (the strand where the DNA polymerase enzyme is travdling in the same direction as
Figure6.6DNAreplic:at""1
115
II Nucleic acids and protein synthesis
the helicase enzyme uncoils the DNA). The copying of the other strand is moving away from the
unwinding site. Here, copying has to be frequently restaned. The resulting fragments in this new
DNA strand have to be joined up. This is done by another enzyme. known as ligase .
DNA polymerase a lso has a role in "proof reading' the new strands. Any 'mistakes' that start
2 Whyaremany to happen (such as the wrong bases attempting to pair up) are immediately corrected so that each
replication forks new DNA double helix is =actly like the original. Then each pair of double strnnds winds up into
required in the a double helix. The outcome is that one strand d each new double helix came from the parent
replication of a single chromosome and one is a newly synthesised strand. This arrangement known as scm.l .conservati n-
strand of DNA? rcplkatk>n bemuse half the original molecule is conserved. We return to this issue shortly.
Crkk aod Watson suggested repjicatioo of ONA woo~ be 'semi-conservative', aod thts M, sir.ce been
shownexperimentally,usingONAofbac:teria 'labelled'witha 'hea~'nitrogenMt<>pe
,faol'f\ti,elynewdooblehel;xwe,e
~~~~x=~·~,u''"
wooldbeconservedwithoot
unzjppingin1henextgenera1Klfl
(i.e . conHrvativareplication)
116
6.1 Structure and replication of DNA I
The confirmation that DNA is replicated semi-conservatively came from an investigmion by
Meselson and Smhl. Jn this experiment, a culture of a bacterium (E,cben'cbia col,; page 2) was
grown in a medium (food source) where the available nitrogen contained only the heavy nitrogen
isotope (nitrogen- IS, "N) for many generations. The DNA of the bacrerium became entirely ' heavy'
as a result. (Isotopes are explaifled in Appendix I, on the CD).
These bacteria were then transferred to a medium of the normal Oight) isocope (nitrogen- 14.
"N). The result was that the new DNA manufactured by the cells was now made d nitrogen-14.
The experimenters then observed and measured the change in COflcentration of nitrogen-15 and
nitrogen-14 in the DNA of succeeding gen.,,-ations. Noce that these obs=tions were possible
becausethebacterialcelldivisioru inaculrureof£. coliarenaturallysynchronised:evcry
6o minutes they all divide. The DNA was extracted from samples of the bacteria from each
succeeding generation and the DNA in each sample was separated.
Separation of DNA was carried out by placing each sample on top of a salt solution of increasing
density in a centrifuge tube. On being centrifuged, the different DNA molecules were carried down
to the k>vel where the salt solution was of the same density. Thus, DNA with 'heavy· nitrogen
ended up nearer the base of the tubes, whereas DNA with 'light' nitrogen stayed near the top of
the tubes. Figure6.8showstheresultsthatwereobrnined.
r
:i
4 ThenalabelledcultureofFm/iwasswitched f
bad lo a medium providing unlabe lled ni trngen te, one gene,ation
000.•• ''NS/OM>ooo,elhe,Sl,w,s )
synchronised.and
· after on egenera tiona lltheDNAwasof
intemiediatedemrty{eachofthedaughte, 11
cellsrnntair>ed(i.e.ron,erved/oneof the
parental DNAstrandsconta ining 1'N alongside
a1>ewlysyn1hes;s..dstrandcontainingDNA afte,lwogene,ations
ma<lehom'4N)
• after two ge ne rations 50% of the DNA was
:
I
I
11
I
11 )
:;;=-~~s::~~.!::~~;~~~::tion.
given thatlabelledDNAwaspresentinonly
haH'thecells(onestrandpercell) increasing density
Figure6.8Experimenttosl>owthatDNA,eplicationissemi-conservative
3 a Pred ict the experimental results you would expect to see if the Me5elsoo--Stahl experiment
{Figure6.8)wascarriedonforth~generatiom
b If replication was by a 'comervative' process, what would be the result of this experiment
afteronegenerationinnitrogen-14( 14N)7
lf)
II Nucleic acids and protein synthesis
118
6.2Proteinsynthesis l
~ad~.:queoo,
~:t::~
thecOOngmand)
,efereocemand
(coding •t rand) cfrectiooooly
the'l'-'fll'lkinfl)(ffi,1\ion
ol!hecodingmand
ils.pecifies!helWll'ill
,eq.,.,oceofaminoacids
thalmakeupapm!ein
Figure 6.9 Part of a gene arid how rt, DNA codes /0< am ino~cids
,,,~ GCT
GCC
Al<]
Arg
phenylalan;ne
"' TTG Am
TCA
TCG
Ser
Ser
m Lys TCT Al<]
proline TTC Lys TCC Al<]
t,yptophan
tyrosine
..'" CGA Ala
CGG Ala
CTA Asp
CTG Asp
CTT Glu
CCA Gly
CCG Gly
CCT Gly
CCCGly
Figun,6.10Theamiooaodsusedinproteir1Sar.dthei,DNAtrl)letcodes
'"
II Nucleic acids and protein synthesis
o o'
tothe3"endofthegrn\\ingRNAmolecu\e
freenucleotidesalignby
nn>>c,c,c,c,--<>" > --<--<n ""' basepaifingandareheld
coding strand
by hydrogen bonds
~
- ~ ~ ~ ~ ~· TTT
fielixne-form,
astheRNA
polymerase
mOYesaloog
120
6.2Proteinsynthesis l
The amino acid is atmched to its transfer RNA by an enzyme. These enzymes, too, are specific
to the particular amino acids (and types of transfer RNA) to be used in protein synthesis. lbe
specificity of the enzymes is a further way d ensuring the code is implemented correctly and the
aminoacidsareusedinthecorrectsequence
ATP provides the energ y for the attachme nt of a mino acid to its specific tRNA.
Eadlaminoaddi<linkedtoaspedlictransle<RNA(tRNA)
before~ can be used inprnt,,;n,ynthes.is. Thi<i<!heprna.•ss
ofaminoac:idactiVatioo.~takesr,laceinthecyl~
~=:::" ~ :,'.7",.,::.r-h
..
t ~
acida A
+ ~
anticodoospecilic
for amino acid,
Figure6.12Aminoacidactivabo n
+ ATP
specific
m,,,,.e
:!=~~~a=:~~;:~UWA,
~
K:3~-
Jn stage three a protein chain is assembled by a proces, called translatio n . This occurs in the
ribosome move,
along themRNA ribo,;o m c (Figure6.13).
You can see that the ribosome consists o f a large and a small subunit. Both paru of the
ribosomeareintimatelyinvolvedin theprocessoftranslation .
• On arrival at the ribosome, the messenger RNA molecule binds to the small subunit at an
attachment site. In this position there are six bases (two codons) d the messenger RNA exposed
tothelargesubunit atanytime.
• lbe first three =posed bases (codon) of messenger RNA are always AUG. A molecule of transfer
RNA with the complementary anticodon UAC forms hydrogen bonds with this codon. The amino
acid methionine is anached to this transfer RNA molecule.
largesubonit • A second transfer RNA molecule bonds w ith the next three bases d the messenger RNA
Figu,..,6.11Ribosomestroctu!1' molecule, bringing a nother amino acid alongside the methionine molecule. Which amino acid
thisisdependsonthesecondcodon ,dcourse
• Whilst the two amino adds are held close together within the ribosome, a peptide bond is
formed between them by condensation reaction. This reacrion is cataly5ed by an enzyme found
inthelargesubunit.Adipeptidehasbeenformecl.
• Now the ribosome moves along the messenger RNA molecule in the 5' ----> 3" direction and the
next codon is read. At the same time, the first transfer RNA molecule (with amicodon UAC)
leaves th ,ibosome, m· us ·1 am·no ad~.
II Nucleic acids and protein synthesis
• Then a third transfer RNA molecule bonds with the next codon of the messenger RNA molecule,
bringing another amino acid alongside the second amino acid residue of the dipeptide.
5 Drawandlabelthe Immediately, a peptide bond is formed between them by a condensation reaction. A tripepcide
structure of a peptide has been formed and starts to emerge from a hole within the large subunit.
bond between two • Again, the ribosome move:s along the messenger RNA molecule and the next codon is read. A!
amino acids the same time, the second transfer RNA molecule leaves the ribosome. minus its amino acid .
• A fourth transfer RNA brings another amino acid to He alongside the third amino acid residue.
Whilst these amino acids are held close together another peptide bond is formed
• By these steps, constantly repeated, a polypeptide is formed and emerges from the large subunit.
Eventually a 'stop' codon is reached. This takes the form of one of three codons - UAA, UAG CO"
UGA . At this poirn the completed polypeptide is released from the ribosome inro the cytoplasm .
The steps of tran slation are shown in Figure 6. 15.
• Free amino acids in the cytoplasm anach ro specific transfer RNA molecules . There is one transfer
RNA molecule for e ach of the twenty different amino acids. The triplet of bases of a transfer RNA
molecule is known as an a ntieodon
• In ribosomes, transfer RNA anticodons align with RNA codons by compleme n tary base
pairing . This brings amino acids side by side and peptide bonds are fCO"med between them.
Apolypepiidemoleculeisformed.
By these steps. the DNA code is translated irno an amifX> add sequence in a polypeptide. There
are different ways of presenting the genetic code and this is the significance of the term "gen e tic
dictionary' . The genetic dictionary is a list of the particular base s,,quences that correspond with
particular amino acids. It depends on whether it is the code in DNA, messenger RNA (or transfer
RNA) thatisquoted.
Ca>l you say why?
Ill
6.2Proteinsynthesis l
0
elongatK1o trar,slatoon
m<~ f~,f
~translaboo ~
~ ,····
)
3
matu,e - ~ S
pmotiooof posrtK>nol
'"'" start codon stop codon
~f~~~~it l
G AA G C UUUU G U U C
la,gesubunitof tRNA-amino
ribosome add complex
withanticockm
{Glua,riving)
two tRNA,
temporarily held
on ribosome
pept.debond
formed between
aminoa6dshekJ
at the ribosome
stop codon
Figu~6.15 T,ar,slation
123
II Nucleic acids and protein synthesis
The DNA triplet codes are recorded in Figure 6.10 whilst those of messenger RNA are listed in
Figure 6.16. These codes do nOI need memorising but their rdationship is important and you do
needtobeablerousethem.
""
""'
aspartic:Mid
cyste;ne ,,
glutam ine
glutamicacid
UAU lyr UGU Cys
glyc;r,e Go UAC lyr UGC Cys
UCA Ser UMStcf> UGA Stop
<KG Se UAG Stop UGG Trp
CGU AA]
CGC Afg
lysine Lys
CCA Pro CGA AA]
CCG Pro CGG Arg
phenylalanine AUU lle
pml;r,e AUC lie MC Asa
MA"'
MG lys
AGA AA]
AGG Arg
GAU Asp GGU Gly
tryptophan
GAC Asp GGC Go
tyrosine GM Glu GGA Gly
GGG Gly
Figure6.16ThemessengerRNAgeneticdictiorwy
124
6.2Prote;nsynthes;, I
,rrm
j.:;';;'""'"'
'''
L.LLL...LL
transcription
1RNA-.,,mlno 1cldcompln
showing complement.Ky
~~-
J
p;,<1ollheresultingpro1Pin~.
~!hearrinoacid~
;,swmbltd:
~ionsloraminoil<m
$hcM/ninF9Jre6.10)
phenoryplcippNr.n:f
ol ......., rfdbloodCl!lsand
sid:lectls(llb'Hb)
125
II Nucleic acids and protein synthesis
An example of a gene mutation occurs in the coodition known as skklc ccll a n.-.cm ia. Here the gene
that codes for the amino acid sequence of a pan of the respiratory pigment haemoglobin of red bkod
ceUsoccasionaUymutate:satoncbasepair(Figure 6.17). Theresu.ltofthischangeisthatthebase
adenine is replaced by thymine at one position along the coding 5trancl. The messenger RNA produced
from this DNA contains the triplet code GUG in place of GAG. This causes the amino acid valine (a
non-polar amino acid) to appear at that point, instead of polar glutamic acid. The presence of non-polar
valineinthebetachainofhaemoglcbingive:sahydrophobicspotintheotherwisebydrophilicouter
section of the protein. This tends to attrnct other haemoglcbin molecules to bind to it.
When a person who is a sickle cell 'carrier' undertakes sudden physical =ercise (c. moves to
a high altitude) the oxygen content of the blood is lowered. Jn these conditions the sickle cell
haemoglobin molecules readily clump together into long fibres. lbese fibres distort the red blood
cells into sickle shapes. In this condition the red blood cells cannot transport oxygen. Also, sickle
cells gee stuck together. blocking smaller capillaries and preventing the circulation of normal red
blood cells. The result is that people with sickle cell trait suffer from anaemia - a condition of
inadequatedelivcryofoxygentocells.
• Nucleic acids are polymers. They are polynudeotides • Transfer RNA (tRNA) molecules occur in the cytoplasm. These
composed of loog dlairn of nucleotide monomers (a pentose moleculeshaveaparticulartripletofba=atoneend(known
mgar,apho:;phatemoleculeandanitrogenoU5base)combined asana nticodon)andapointofattachmentforaspecific
together. There are two forms of nucleic acid, DNA and RNA am inoacidattheotherend. TheroleoftransferRNAisto
TheseformsdifferinthesugarandinthebaSe'itheycontain, pickupfreeaminoacidsandbringthemtothemessengerRNA
and in their ro les in the cell. ONA is found in the chromoso= intheribosome.Complementarybasepairingbetweenthe
and RNA in both the nucleus and the cytoplasm. ONA also codons of messenger RNA and the anticodons of transfer RNA
ocaminmitochoodriaandchloroplasts results in the original ONA code being translated into an amino
e ATP istheunivers.alenergycurrencymoleculebywhich acidsequenceintheprocessofpo lypept id ea nd protein
energyistransferredtodousefulwork.ATPisasoluble synthes is.
molecule, formed in the mitochondria but able to move e lhereare threestagestopolypeptideandproteinsynthesis.
intothecytomlbyfacilitateddiffusion.ltdiffusesfreely - lnthenucleus,acopyofthegeneticcodeofageneis
aboutcell1 made in the form of a single strand of messe nger RNA in
e lhesequen ceoffourbases,adenine(A),thyminem, the stage called transcription. ThemessengerRNApasses
guanine {G} and cytosine (0, along a DNA molecule is the out into the cytoplasm
geneticcode.Thisisathree-lettercode - threeofthefour - lnthecytoplasma minoacid activation occurs.Eachof
basescodeforaparticularaminoacid. The sequence of bases the 20 amino acids is attad.ed to a specific trans fe r RNA
in the DNA molecule may be transcribed to a complementary
sequence in messe nger RNA (mRNA). Once formed, - =""'"
Finally a new polypeptid e is assembled when the
messenger RNA pasSe'i out of the nucleus into the cytoplasm informationofthemessengerRNAis trans lated ('read')
and to a ri bosome where polypeptid e sy nth es is occur1 in a ribosome and the transfer RNA molecules with their
DNA 'codes' are referred to as triple t codes and those in attac:hedaminoac:idsarealignedintheordergiveninthe
messenger RNA are called codons code. Peptide bonds are formed between the amino acids
126
Examination style questions I
Examination style questions
1 Fig. 1.1 shows the replication of one strand of a ONA double 3 A molecule of messenger RNA {mRNA) was produced during
helix. thetranscriptionofageoe.Partofthetemplatesequenceof
ONAwasATGC
The diagram shows the part of the molecule of messenger
ox
RNAcorre;pondingtothatsequenceoffourbases
~LclR,::c~
~ HOH H,f,{"K\ E
H H N_).. \
? OH H~Jl.)'.~H
o-1 o-----<:~'o
O H H H,t,/"'H
H H H--..,)..,
OH JL ..1
0-~p~
F'<Hf 0 0
_;JN,H~
" ' "'";".
a) Name W to Y [3)
b) Explain how the structure of ONA enables it to replicate
o-e---0-clR2
6 " "
0 A
semi-comel'Vatively. [3]
" "
? °"
c) Explain why it is important that an exact copy of ONA i1
madeduringreplic.ation [2] a) Name the parts of the mRNA molecule shown in the
[Total: 8] diagram above labelled D, E, F and G (4)
(Cambridge /ntemationa/ AS ;md A Level Biology 9700, b) Copy and complete the t able to show threewa~ in
Paper 02 QI June 2005) which mRNA differs from ONA. [3)
127
AS Level
7 Transport in plants
Flowe ring pla nts do not have compact bod ies like th ose of a nima ls. where they a re a bsorbe d or produced to w he re th ey ue stored
Leavesandntensiveroot systemssp read o uttoobta inthe lig ht o ru sed .Pl a nts do noth avesyste msfortra nspo rting o xyge n a nd
e ne rgy.wat er,mine ra lio nsa nd ca rbo ndioxide tha tpla ntsga in carbondioxide; inst eadth ese gases diffuse through a ir spaces
fro m th e ir e nvironm e nt to ma ke o rga ni c molecul es, suc h as s ugars w ithin s te ms, roots a nd leaves.
a nd a mi no acids. Tra nsport systems in pl a nts move s ubst a nces from
Plants have two transport By the end of t his section you should be able to:
tissue~: xylem and phloem
a) draw and label from pre pared slides plan diagrams of transverse sections of ste ms, roots and
INVes of herbace ous dicotyledonous plants using an e~piece graticule to show tissue in correct
proportions
b) drawandlabelfrompre pared,lid.,. thecellsinthe diffe rl!n tti«ue inroots,ste m,andleave,of
he rb..u,ousdkotyle donou,plantousingtranwerseandlongit udinal,«tion,
c) draw and label from prepared did"" the structure of xyle m v=e l e le me nts, phloem ,ieve tube
e le me nts and companion ce ll, and b e abl e to recognise th""e using the light microscope
d) relate the structure of xylem vesel e lemenu, phloe m ,i""" tube e le ments and companion cello to
128
7.1Structureoftransporttissues .
The stem
The role of the stem i., ro support !he leaves in the sunlight and ro transport organic materials (mainly
sugar and amiflo acids), ions and ,varer between the roots and leaves. Thi., tr:mspmt occurs in the
mscular bundles. The vascular bundles of the stem occur in a ring ju SI below the epidermi.s. To
theoutsideofthevasrularbundle.slicsthe=it=andtotheiruideisthepith.Bo!hconexandpith
mostly=11si.stofliving,relativdyun.specialisedcellswiththincellulooewalls(Figure7.6, page!J3).
youngsunflowu plant
:;!,'~:':~t:'~J0jcu1open10 !heb<anchin,g
vasrularsy,temol!hestem vascu!arbundlP<
capolfibrP< - loog,Mrrow,
tapefingcellswith!hickened
wallsmengthened'M!hlign;,,
Thesl'cellsa"-'!'DNwilhoutli,,;ng
coments.Rb«>s,wilh!hefl'St oi
1nevascularbundle,pro,,;c1epan
ofth,eml'Chanicalsuppo<1lorthe
aerials)"lem(stem,Jeaves,flcJM>")
camb<,Um(cels
able!O<ivide)
~:':'u~~buo;:"'!ff~~
majnroleisth,epassageof
waterupthl'plant
photomicrographofsunflow ersteminTS(x30)
Figure7.1Thedistributionandstructureofthevasru!arbundlesoithl'stem
129
II Transportinplants
The leaves
The leaf is an organ specialised for phoiosymhcsis the process in which light energy is used
to m3ke sug:,r. Carbon dioxide from the 3if and wmer from the soil are the raw materials used,
oxygenisthewasteproduct.
The le3f consists of 3 le3f blade conneaed to the stem by a leaf stalk. The blade is a thin
structure with a large surface area. Within the leaf blade 3re mesophy ll cells th3t are packed with
chloroplasL~ - the org:melles in which photosynthesis occurs. Th'""' is a thin film of water on the
outer surface of these cells. lbe e pidermis is a rough, transparent, single byer of cells surrounding
the me.sophyll tissue. There is an =tem3l waxy c uticle to the epidermis which is impervious. This
reduceswatervapmu-lossfromtheleafsurface(Figure7.2).
The epidennis has m3ny tiny pores. called stomata - the sites of gas exch3nge. Within the leaf's
mesophyll tissue are continuous air spaces. Throughout the leaf bbde is a network of ,.,._..cular
bundles. These provide mech3nical support but the le3f is also supported by the turgidity of 3l[ its
cells. surrounded as they are by the tough epidermis. Leaves are relatively delicate strucruresand.
attimes,theyhavetowithstanddestructiveforcessuchaswindandrain
palisade mesophyllcell.
chief,;teofphotosynthes;s
;nthegreenplant
,,.::::o· <.ytoplasm (J
vein.rntin
LSandTS
8
0
Figure7.2The,tructuresoftheleal
130
7.1Structureoftransporttissues .
The root
The root anchors the plant in the ground and is the site of absorption of water and ions from
the soil. The main rnp roo1 fonns bternl roois, and growth continues at each roo1 tip. The growing
plant is continuously making conma with fresh soil. This is imporrnm because the thin layer of
dilute soil solution found around soil panides is where the plant obtains the huge volume of water
it requires. Minerals salts as ions are also absorbed from here.
The vascular tissue of the root occurs in a single, central s i de . Around the stele is a single
1 Outlinetheproperties
byer of cells, the e ndod~·rmls. These cells have a waxy strip in their radial walls, known as
of cellulose that make
the Cas p a ria n st ri p . Immediately within the endodermis is another single layer of living cells,
known as the pericycle (Figure 7.J). Only at the base of the stem is the vascular tissue of the
root reorganised into several vascular bundles arranged around the outside of the stem.
Sea Rg7.8
,-.
reglooofgru.vth
(cel<!Msklo)at
Figure7.3Thestructureoftherool
2 The external dimemiom, volume and surface area of six geometrically shaped objects -
threeofacompactshapeandthreethatareflatandthin - arelistedinthetablebelow.
Compact
Luge
Dimensions/mm ""'"
1 x 1 x 1 l x l x l 4x4x4 lx 1 x0.5 8x l x0.5 16x8x0.5
Volume/mm•
Surfaceuea/mm• 6
a Cakulate the surface area to volume ratio of the c Comment on the effect of shape on the surface area to volume ratio of
six objects anobjectasthesizeincreases
b Comment on the effect of shape on the !.Urface d If theo;e objects were organisms, what are the implications of their size
area of an object as the size of the object increa1es and shape on the movement of nutrients or waste products by diffusion!
131
I I Transportinplants
xylempa,ench)mil
(!heonly i,,;ng
cellsofx;,<'m
tissue)
ligninthiden;n9as,pira!s.ring,.netw<rl:
0<soldblocks - depo,itedooinsideof
Figure7.4The,tructureolxylemtissue vessel.,uengthening!heceluloselayer,
During dt,-velopment, the end walls are dissolved away so that mature xylem vessels
are lo n g, h o llow m bes. The living contents of a developing xylem vessel are used
upintheprocessofdepasitingcellulosethickeningtotheinsideofthelateralwallsof
the vessel. This is hardened by the depo,;ition of a chemical substan=, Hgnln. Llgnin
is a substance that makes cross links with the hemicelluloses that fill the gaps becw=n
cellulose fibres. This greatly strengihens the =llulose and makes it impermeable
to water. Consequently. xylem is an extremely tough tissue. Furthermore. it is
strengthened internally, which means it is able to resisl negative pressure (suction)
without collapsing in on itself. When you examine xylem vessels in longitudinal
seaionsbylightmkroscopyyouwillseethat theymayhavediffcrentlydepasited
thickening; many have ring.s of thickening, for example. O!her xylem vessels have
more massive thickenings. However, all xylem vessels have areas in their lateral walls
(includingthepitswherethe=llulo.sewallisespeciallythin)inwhichthereisalayer
of cellulose but no lignin. Here the walls are entirely permeable and permit lateral
mO'l'ements of water to surrounding tissues. This is role of the xylem - ro supply
watertothelivingcellsoftheplant.
Flbresalsooccurbecw=nxylemvessels,therebysirengiheningthewholebundle.
Interestingly, in ferns and cone-bearing tr=s, xylem tissue is absent. Here the water-
conducting tissue consisls of fibre-like lrach d d s with large pits in their lateral walls
throughwhichwaterpassesfromtracheidtotracheid(Figure7.4).
fibresareathod-walledtissue.wilhwal!sof
cellulose,tre<>gthenedbyl>gn;n - whenmature
thecellcontentsdie;fib,._.,a,elong.narmw.
pointed.emptycel!swithpitsintfieirwall,
wherethelivingcontent,onceconnected
Figure7.5The,tructun,offib,._.,
Ill
7.1Structureoftransporttissues .
Phloem tissue consists of sic,.., mbcs and companion c ells, and is served by transfer cells in
the leaves. Sieve rubes are narrow, elongated elements, connected end to end to form rubes. The
end walls, known as sieve plates, are pcrfora!ed by pores. The cytoplasm of a mature sieve tube
has no nucleus, nor many of the oiher org:mdle.s of a cell. However, each sieve tube is connected
to a companion cell by strands of cytoplasm called plasmodesmata, that pass through narrow gaps
(called pits) in the walk lbe companiOfl cells service and maintain the cytoplasm of the sieve
tube, which has Jost its nucleus. Phloem is a living tis.sue, and has a relatively high rate of aerobic
respiration during transport. In fact, cranspco1 of manufactured food in the phloem is an active
process,usingenergyfrommernbolism(Figure7.6).
photomOClographofLSphkieminHP{x200)
phloemisatissuethatmaybecomecrushed
anddam;,gedasstemtiss.ueissectKlnedfor
cytop!asmKcoonecbons
betwe<'ncompanK>ncells
and sieve tube contents
cytop!..smloses nudeus
andmanyotganelles
phk>emtubesarethes iteofmovementof
suga,aOOaminoacid,
Figure 7.6 Tfie,tructureofphloemtie,;ue
133
II Transportinplants
With guidance. the different tissues a slide shows can be identified and their precise distribution
represented in a pla n d iagram (also called a tissue map). This will show their relative positions
to scale. Individual cells are not shown. Then the size of the specimen can be calculated and the
3 Orawandlabela magnification represented by a scale bar.
plandiagramofa Figure 7.1 has the necessary information to create a plan diagram of pan of a plant stem.
representative part of the However, it is imponant you create your own plan diagrams by =amining prepared slides, using
sectionofthe1temin an eyepiece graticule (page 9) to show tissues in correct proportions. Follow the guidance on
Figure7.1toaccurately recording observations in drawings on page 8.
record the distribution of Figure 7.2 has the necessary information to create a plan diagram of pan of a plant leaf.
theti1sues However, it is imponam you create your own plan diagrams by examining prepared slides, using
Using the magnification
an eyepiece graticule to show tissues in correct proportions.
datagiven,addan
Figure 7.3 has the necessary information to create a plan diagram of pan of a plant root.
appropriate scale bar
However, it is imponam you create your o wn plan diagrams by examining prepared slides, using
line to your drawing
an eyepiece graticule to show tissues in correct proportions.
134
7.2 Transpon mechanisms I
4' 7.2 Transport mechanisms
Movement of xylem sap By the end of this section you should be able to:
andphloemsapisby
mass flow. Movement in a) explain the moveme nt o f watf'r betwe..n plant cf'lls, and betwf'en thf'm and th e ir f'nvironment,
thexylemispassiveasit intf'muofwatf'rpote ntial
t5drivenbyevaporation b) explain how hydrog en bonding of wate r molecul es is involv..d with movf'me nt in th e xyle m by
fromtheleaves;plant:suse cohesion·tensionintranspirationpullandadhesionto,e llulose cf'llwalls
energytomovesubstances c) describe the pathways and expl a in the mf'cha nisms bywhkh water and minera l ions are
in the phloem tran,pone d from sc,il to xyle m and from roots to leav..,
d) df'finethe te rmtranspirationand explainthatiti,aninf'Vitable conse quf'ncf' ofgu ex,hange
Xylem sap move1 in one
directionfromtherootsto in plants
therestoftheplant.The e ) investigateexperimf'ntallyandexplainthe factor,thataffe cttranspira tionrate usingsimpl e
phloernsapinaphloem potomf'tf'rs,leafimpressions, e pide rmalpef'ls,andgrid,ford et er mining,urfacf' area
sieve tube moves in one fl makf' a nnotated drawing,, using pre parf'd slides of cross-sections, to ,how how leav..s of
direction from the location xerophytkplanuarf' adaptedtorf'du,e wate rlossbytran<piration
whereitismadetothe g) state thatauimilatf's,suchassucrosea ndaminoacids,move bf'twef'nsources (e.g.le avf's
locationwhereiti1used andstora gf' organs)andsink<(e.g.bud,,flowers, fruits, root,andstoragf' organs)inphloe m
orstored.Atanyonetime sie ve tubes
phloemsaprnnbemoving h) expl a in how sucrose is loadf'd into phloem sif'Ve tubes by companion ce lls using proton pumping
indifferent directions in and the co4ransportf'r me chanism in the ir ce ll ,urface mf'mbranes
different sieve tubes i) explain mass flow in phloem ,ap down a hydrostatic pressure gradie nt from source to sink
13S
I I Transportinplants
Figure7.8Roothai1' thesiteofabsotpbonanduptake
The very large numbers of delicate root hairs form behind each ro« tip (Figure 7.J). These are
in close contact with the soil solution which has a very high water potential. (Remember, water
potential, the name we give to the tendency of water molecules to enter or leave solutions, wa.s
introduced in Topic 4, page 83.) At the same time the cells of the root cortex have a water potential
much lower than that of the soil solution. Water emers the root from the soil solution, passing
downawaterpotentialgradient(Figure7.9:,).
Anotherprocessoccurringintheroothairsistheactive uptakcof solublcionsfrornthesoil
solution. These ions include nitrate ions, magnesium ions and several others, all essential to the
metabolism of the plant. Once absorbed. the ions are transported across the cort= of the root.
4 What features of
They converge on the central s!ele and there they are actively secreted into the water column
root hairs facilitate
Oowing up the xylem. The continuing movement of ions across the cortex of the root is partly
absorption from the
soil? responsible for the gradient in water potemial between soil solution and the cells of the stele. We
returntotheissueofionuptakelaterinthisropic.
136
7.2 Transpon mechanisms I
., From root hairs to xylem - apoplast and symplast
Look at Figure 7.9b. It shows the two pathways that water can mke through plant tissues.
The alternative pa1hways of waler mo,..,mem shown are:
• the apoplast. This is the interconnected 'free spaces' that occur between the cellulose fibres in
the plant cell walls. These free spaces make up about SO per cent of the volume of the cell wall.
lbe apoplast also includes the water-filled spaces of dead cells and of the xylem vessels. Nore
thmthisrouteentirelyavoidsthelivingcontentsofcells.Weshallseethatmostofthewarer
passing through plant tissue travels by this pathv,ay, mos1 of the time - a case of transport by
• the ,.-ympla,;.1. This is the living content5 of cdls. W..ter enters cells through the cdl surface membrane
by osmosis and diffuses through the cytoplasm of cells and the cytoplasmic connections between cells
(rnlled plasmodcsm:,la). You wm remember that the cells are packed with many organelles. These
tend to slow the flow of water, so this pathway is nOI the major one for water flXNemem.
So, most water travels across the roo1: via the apopbst pathways. until that is, the endodermis, is
b}
Figure7.9Wate,movement-fmceandpathways
a)Thewate,potential(o,jg,adientofthewholeplant
b)Thewatl'l'polential{.,)gradientaacs,celkoftf>emotcortex
137
II Transportinplants
J?to
~--
A";.~.::""'""
l'Yapo<atboofwater,aod
tl uttimately"-"poo~ble
forclrawingwaterup!he
piam,tem{!he
transpifatioorueam).
-~
- 1: · . ••,.,... . ,~"-
waterpa<ses!hroogh
l ; C)'toplasm(apoplasl:
: . .• • • pathwaybbded,
----- - ~ teq,orarily)
-,~',,
,:=,
between cell,
(plasmodesmata)
Figure7.10Wateruptakeandlos,bytheg,,...np!ant
138
7.2 Transpon mechanisms I
Transpiration and the role of stomata
Transpiration :theprocess
Transpiration is the name given to the Joss of water vapour from the aerial parts of plants. lbe
through which water
vapourislostfromthe mesophyll cells in the leaf are moist and their walls contain water between all the cellulose fibres
aerialpartsofplants.11 there. Consequently, there is a film of water around the outside of the cell walls, too. So water is
occursastheresultof freely available to evaporate from the cell walls and it does so. The air inside the leaf is usually
evaporation of water at saturated with water vapour.
thernrfaceofmesophyll Stomata are pores in the epidermis of leaves. Each stoma consists of two elongated guard
cells into the airspaces cclls. lbeseareattachn:ltothesu1TOUndingepidermalcellsandjoinedrogetherateachendbut
withintheleaf,followedby they are free to separate to form a pore between them (Figure 7.11). The epidermal cell besides a
diffusion of water vapour guardcelliscalledasubsidiarycell.
outoftheleaf,mainly The water vapour that accumulates in the air spaces between the mesophyll cells diffuses OU!
throughstomata,downa
through the pores of open stomam, prmided there is a concentration gradient between the leaf
water potential gradient
spaces and the air outside the leaf. This difference in concentration of water vapour between the
from the surface of
spongymesophyllcellsvia leaf and the =temal atmollphere is highest in hot, dry. windy weather. Transpiration is greatest in
airspacesintheleaftothe these conditions. as we shall shortly demonstrate. The site of transpiration is shown in Figure 7.12.
atmosphere
subsidiary cel l
associatedwithguan:l
cell (conta insno
chloroplasts)
cell contents have
hardened and shrunk
dur;ngfixationof
Jeaf tissue
thockoute,wa ll
ofepidem,is
.....; J - " ' ! -- co"",ed byawaq
cutode
~~~_ _ L _ _ _ _ _ porepartially
guan:lcell but dorsal and epidemial
hasth;n ...,nt,alwalls are cells
lateralwalls greatlythockened
withcelulose
Figun,7. 11Thestructure ofstomata
139
II Transportinplants
Figure7.12Thesiteoftranspiratioo
sandwichthepink"standard"betweenstripsofstickytape
~ -j· t''
cut ml
Figure7.13Cobaltchkiridepape,llSl'dtomeasurewatervapourlossfromaleafsu,lace
140
7.2 Transpon mechanisms
watercanbedrawnuptoagreatheightwithoutthecolumobfeakingoff
I
Dfpu llinga?,Jrt
Investigating transpiration
The conditions that affect transpiration may be studied using a potomctcr. You will find that these
are also the conditions which effect evaporation of water from any moist surface. They include the
humidity of the air, temperature and air movements . In Figure 7. 16 the ways in which these factors
maybeinvestigatedissuggested.
Actually, the potometer measures water uptake by the shoot. In the potcmeter the shoot has an
Fig ure 7 .1 S Xylem,ve,selwith unlimited supply dwater. By contrast, in the intact plant the supply dwater from the roots may
internal ring, of lignin thickening slow down (as in drought conditions, for example), so the water supply is also a factor in the rare d
their lateral walls !rnnspiration in the intact plant. This point will help you answer Question 7.
141
II Transportinplants
Table7 .1 ~actorsthatinftuencetrar,sp;,aticmandhowtoinvestigatethem
Thti facto" that inf luenat tranopi ration: How to inn stigate facto r using the potomet..-:
humidity - at km humidity m0<e water evaporates compare transpirabon by shoot contained in
polyther,ebagwiththatbyshootinmovingai,
tem perature-heatenergydrivesevaporation from compa,e trampirationatdifferentai,
rn,faceofmesophyUcells temperatures
a·rmovement - ·ndcarf awaysaturateda'r c°""'aretranspi,ationatdifferentfanspeeds
fromaroundH'aves.maintainingaconcentration
gradientbetween!ealinteriorandair outsidestomata
water suppl y- if !eaves become fliKdd the (not applic:abH' - potomete, supp l>es unlimited
stomata close water to shoot)
Thepotomete,maybesetup ul>de,differen!en,,;,onmental
conditions(e.g. l'9htv. dark;lowv.hql tefTI!)l'lature;mow>gv. sti ll
!ealyIDOOl(lose'.I at ). and the rateoluanspirationmeasu red
7 Explainthe5ignifi(ance water""!)OOrtoair)
ofthefactthatinlarge
treesthereisaneas ily
detectable shrinkage water drawn up the
stemto,eplac~ the (thecaplllarytubeisrl'Charged
in the diameter of
watertraospi,l'd withwaterfromhe,e)
the trunk during the
daytime inhot111eather
tap(closed) in a given ti me
that recovers in the
night
capillary tube
(1mmind00\e!er) aswaterisdrawnintotheJ)lam
amenisru,;appea"hereand
moYeSaloogtl>ecapiltarywbe
" ' = = ~~ ~
Figure 7. 161nvesbgatingtrarispirationusingapotometer
142
7.2 T,anspon mechanisms I
On the other hand, some plants are wdl able to survive and often grow well in habirnts where
water is permanently scarce. lbese plants are known as x erophytes. 1bey show fe:itures
that directly or indirectly hdp ro reduce the water loss due to transpiration to a minimum .
The adaptation., they have thm enable them to survive where water is short are referred to as
xero morphic fea tu res. lbese are summarised in Table 7.2.
Table7.2Ti,,kalstructu,alleatu,esolxemphytes
Exceptionally thH cuticle to ~af (ar,d stem) Prevents wale, loss thmugh the external wall of the epidermal
epodermi, cells
Llyerofhai"ontheep,de,m!, T,apsmoistairo,erthe~afandreducesdiffusion
Reduction in the number of stomata Reduces the outlets through which moist a i, can diffuse
Stomata in pits or gro,es Moist air is trapped outside the stomata, reducing diffusion
Lea f rolledmfokJed \\+.en,hortofwate, Reduces area ffom\\+.khtrar,spi,atk>ncanocrn,
(ce!l,Aac:c,dJ
8 Suggestwhyitisthat,
of all the environmental Leafareasev..,.elyreduced - possiblyto Stemstakeo,eraphotosynthetic:functionandsupportmuch
fac:tonwhichaffect 'needles' reduced leaves
plant growth, the is!.Ue H~ve t\\'o types of mots Obtainwat..,.byboth
ofwater~pp!yis'iO a)superficialroot, expk,itingo,..,.nightcondensatk>natsoilsurface
critical b)deepafldextensiveroot, expk,itingadeepwatertableintheso il
Sorghum, a grninplamthatthrivesinextremelyaridconditiOfls,showsrnarkedadaprntionsinits
leaf structure and roo1 sy.s1em. Marram grass is a pioneer plant of sand dunes.
Examine Figures 7. 1 7 and 7.18carefally. Wb icbif tbefe<Uureslisted in Table 7.2d-Oeseacbplant
exhibit?
I r h. ,~,.,~...............,,,,"""'"'"'"m"
~ ~· ,~·~;,;;:
.
::."Xt~
,klwswate,
•• ~~.'.,.,~,!';
• H',lf.,;am,mata
Figure7.17So,ghum,adrought-resistantlrcpicalplantoleconomK;mportance
143
II Transportinplants
TSofl eaf(~SO)
~~~~~!:ing,rast
Figure7.18Marrangrass/Ammophi/d),apartolsanddunes
144
7.2 Transpon mechanisms I
Signs and symptoms of mineral deficiency
In the soil, the ions dissolvn:l in the soil solution come from the mineral .skdeton of the soil by
chemical erosion . They also come from the decay of the dead organic matter that is constantly
being added to the soil. Among these are nitrate and magnesium ions , lxxh required for 10 \Vhyandhowi:sthe
pankular biochemical pathways (fable 7.J). Consequently. plants tha! are deficient in these supplyofessentkllion5
(and other) essential nutrients normally show characteristic signs and symptoms. Nocice that the maintained in the soil
inagriculture7
signs of deficiency of nitrate and magnesium ions are very similar.
Can you think why this is? Remember, the green colour of pbrus is due to the presence
of chlorophyll . Are nitrate and magnesium ions required for synthesis of new chlorophyll
molecules?
When deficiency sympcoms appear in cultivated commercial crops, the mineral nutrients
avaibble in the soil may be checked by chemical analysis, to confirm the deficiency. lben
anificial fertiliser can be applied to counter the deficiency and restore normal grov,th patterns.
Table 7.3 Nitrate and magr,es;umc symptom, of deficieocy and roles ;n metabolism
Symptoms of deficiency
Nitrate de ficiency Key ingre di en tin amino acid(andprotein} synthesis
upperlea,,...lightgreen.
'°"""'le"""')"'llow
,.,~,..,..,.,
redtKir>g power (NADH)-----...
j
NO a
frnm,espiratkm
2 Aromon;um km, are combined with an o,ganic acid to fDflll the amino a<lds required for
protein synthesis
~\
crganKac:ids amino acids
3 fmmthepoololaminoac:idsarefom....Jthepolypeptidesandproteinsthecell ,equ;,es
r,:::,~;:;;::;::."
is s y n t h e s i s e d ~ . ~"" l
~~
=--~ loonatoooof am<•"
amino acids
peptide bonds COOH
Nudeicac:id,a!socootainnitmgen
(continued)
145
II Transportinplants
(cOfltinued)
Symptoms of deficiency
M agnesium deficiency
1ow,,,1e,.,...ye11ow
fmmma rgin<irrwards,
VPln<fl'fllamge'-"l
CH '"''."'"'"'""'.'
t1d>loroph~ I a
lomph~ I bhasan
hydegoop(---<:HO)
theplaceof theCH,
CH, porphj<'1ring
(l,t,lahsorDing
o o o~ ,,
0
'
}--.~1=~"
ahydrocarboo1aW
~ rn,
Translocation
Tr:mslocation is the process by which manufaaured food (sugars and amino adds
called assimilates) are moved arou nd the plant. This occurs in the phloem tiss ue of the vascular
bundles (Figure 7.1). 1be sugars made in the leaves in the light are moved around the pbnt as
sucrose. For example, youn g leaves =pon sugars to sites where new stems, new leaves or new
roots are being formed. Then, in older plants, sucrose is sent to sites of storage. such as the o:irtex
ofrootsorstems,andtoseedsandfruits.
Meanwhile, nitrntes are absorbed from the soil solution by the root hairs. Nitrates are one of the
raw materials used in the synthesis of amino acids and also nucleic acids. Amino add synthesis
rnkes place in the roots. Amino adds are then moved to sites in the plarn where pro1:ein synthesis is
occurring, such as in buds. young leaves, young roots a nd developing fruits. So you ran see that the
contents of phloem (known as sap) may vary. Also, it tru1y flow in either direction in the phloem.
146
7.2 Transpon mechanisms I
compa nion ce ll a nd s ieve tube
e lement in LS (high power) Seve pl ate ·n surface v ·ew
s.ievettl,eelementwith
erdwak peifo<atedas
a sieve plate
lining!ayerofcytop~m
withsmal mitochoodfia
and some endoplasmic:
reticulum.but without
nudeus,ribmomes0<
p!asmodesmata - Golgi awaratus
cytoplasmic
coonKtionswilh
sieve tube
cell cytoplasm
Fig..-e7.19Theflnestructu,eolphloem
Phloem t ransport
Phloem transport occurs by m ass flow but the pressure difference that drives it is produced by a
11 Whatdoesthe differeru mechanism from thm in xylem transport. Most significantly, translocation requires living
preseoc:eofa
cells (Figure 7.20). Cell respiration is the process by which energy is transferred to drive me1abolic
large number of
re3ctions 3nd processes. Much of the respiration occurs in the mitochondria. ATP is the molecule
mitochondria in the
produced in mitochondria to transfer energy. We 5h3Jl see th3t ATP is involved in the mech3nism of
companion cells
rnggesttoyouabout phloem transport .
their role in the Tran.slOC3tion rnn be illustrated by the movement of sug3r from the leaves. lhe sro,y 5tarts 3t the
movement of 1<1p in poim where sugars are made and accumulate within the mesophyll in the leaf. This is the source area.
the phloem? Sugars are loaded into the phloem sieve tubes through transfer cells (Figure 7.21). From here,
12 ExamineFigure7.20 sucrose is pumped into the ccmpanion cells by the ccmbined action of primary and seconda ry
carefully. pump,;;. These pumps are special proteins in the cell surface membrane. The primary pumps
a Whato;equeoc:eof remove hydrogen ions (protons) from the cytopl3Sm of the ccmpanion cell, so setting up a gradient
events would you in concentration of hydrogen ions between the exterior and interior of the companion cell. This
anticipate in a leaf movement requires ATP. Hydrogen ions then flow back into the companion cell down their
stalk as the content
concentration gradient. This occurs at specific sites called secoo,bry pumps where their flow is
ofawater-jac:keti1
linked to the transport of sucrose molecule., in the same direction.
raised to SO"C?
A!J sucrose solution accumulates in the companion cells it moves on by diffusion into the sk>ve
b Howwouldyou
tubes, passing along the plasmodesmata (Figure 7.19). The accumulation of sugar in the phloem
expect the phloem
rnpsampledfrom tissue lowers the W3ter po1enti3l 3nd wa!er follows the sucrose, diffusing down a wa!er po1ential
a sieve tube near gradient. Thiscreatesahighhydrostaticpressureinthesieverubesofthesourcearea.
leaves in the light Meanwhile, in living cells elsewhere in the pl3nt - of!en. but not necess3rily in the roots,
andatthebaseof sucrose may be converted into insoluble S!arch deposits. This is a s ink area . A!J sucrose flows
the same stem to out of the sieve tubes here, the water potential is raised. Water then diffuses out down a water
differ? po1ential gradiem and the hydrostatic pressure is lowered . These processes create the difference in
hydrostatic pressures in the source and sink are3S that drives mass flow in the phloem.
147
I I Transportinplants
cooc:lusion:livingcellsare
Note:inneither experimentdidthe PSsentialfmtranslocation
leafbla<lewilt - xylemtrensportis
ootheatsensitivea t thisrangeof (b) at3"C. compared with 20"(
temperatu,es(becausexylemvessels tfar,slocatk>oolsuga,fromleaf
are dead. empty tubes) bladewasreducedbyalmmt10%
ofleafd,yweightove,agiventime
conclusion: rateofmetabrnic:
ac:tivityofphkiemcelisaffects
rateoftramlocation
Ffgu,.., 7.21 Transfer cells and the TEM of a leaf vein showing sieva tube alammts. transfe r calls, xylem VRHsels and masophyll cells (x1500)
load ir,gofsievettbes
xylemvasHls
with thickened,
i!1'ifledwals
andpermeatile
p<t il<l'ilS- here
wate,canpass
::•uffOUnding
148
7.2 Transpon mechanisms I
Table7.4Evidencelortr.ep,essureflowhypothesis
13 Vv'hat conditiom
Thecontentsof thesievetubesareuriderpressureand Phloemtissuecaniesmanufactu!l'dfoodtovarioos
maintain
sugarsollil,Onexudesifphkiemiscut destinations(ind ifferent,;evetubes),ratherthanto
a thehighwater
the greatest sink.
potential of the cell
oftherootcortex A.,propriategradientsbet...-een'sou,ce'and'sink' SH'veplatesareabarrie,tomassflowandm>ghtbe
tissuedoexist expectedtohavebeen'lost'in tfiecourn,o/evolution
b thelowwater
ifm.,..flowisthemechanismof transport
potential of the
mesophyllcellsofa
greenleaf7
~ti!,,,~k!~O::~'in~~
,,,,_
concemratK>llinAandBis
waterftowby
hyd=taticp<l'SSUfe
How of water
(=,ylem)
high~drostaticpres:sure
duetods'°""'dsugar
mass flow
along sieve tube
;::=ufromhigh
hydrostalic
Klwr,ydmstati::pn,ssu,e
~!:~~:.~~;.,
wateruptaKeinrootllair
sinkcel ,e.g.starrh
,m,agecell(=B)
Figure7.22Thepressu,ello\\·hypothesisofphloerntrallSport
149
I I Transportinplants
• Internal trilllsport within plants occurs by mass fl ow but there thecc hesiveandadh es ivepropertiesofwater,andmakes
i5 no pumping organ. Two separate tiswes are involved in po5sibletheupwardtransportofwaterinthexylemby
transport.Waterandionstravelinthexyle m,asystemofvessels cohesion-tension.
coonectedendtoeodtoformnon-livingtubes.Manufactured • Transportofmanufacturedfoodinthephloemisby a ctive
foodsarecarriedinalivingtissue,the phl oe m,coosistingof tra nsport requiring living phloem cells. According to the
sieve tubes and companion cells. The xylem and phloem make mass fl ew hypoth esis, solutes flow through the phloem
upthevasc ularti ssuethat branchesthroughouttheplantbody fromaregionofhighhydrostaticpressuretoaregionoflow
andservesroots,stems,leavesandgrowingpoints(buds) hydrostaticpressure.Hydrostaticpressureishighincellswhere
• Watermovesfromthe50iltotheaerialsystembyagra die nt sugarisformed,thesourcea rea,butlow in sink a reas,where
in wa te r po tential. Water is drawn up the stem by a force sugari1convertedtostarch
generatedintheleavesbytheevaporationofwater vapour • Plantsthatsurvivepermanentlyaridccnditions{xerophytes),
fromtheaerialsystem(tra nspir atio n} includingthecropplantSorghumandthesand-duneplant
• Hydrogenbondingbetweenwatermolecules, and between Marramgrass, haveleavesadaptedtoreduc:ewaterlo1sby
the1emolecul~andhydrophilic1urfaces,i1responsiblefor transpiration
Q acellwallstrengthenedwithlignin
Fig.1.1
R atissuethattransporuassimilates [3[
b) Name two assimilates that move from the palisade b)Vasculartissueinrootsissurroundedbyanendodermis
mesophyllcellstothevasculartissuetobeexported Describe thefuncticnoftheendodermisin root~ [3]
c) Stateandexplaintwoway1inwhichthestructureof
from the leaf [2]
c) Explain, using the term w ate r pote nti a l, how water aphloemsievetubeisadaptedforthetransportof
assimilates [4]
movesfromthevasculartissuetotheatmosphere. [4]
[Total: 10)
[Total: 12)
(Cambridge International AS and A Level Biology 9700,
(Cambridge International AS and A Level Biology 9700,
Paper 22 Q5 June 2009)
Piiper 02 Q2 June 2005)
15()
AS Level
8 Transport in mammals
As a nima ls become la rge r, mo re complex a nd mo re active, tra nsport re qui re much g reater s uppli es o f oxyge n. This is tra nsported by
syste ms become essentia l to supply nutrie nts t o, a nd remove waste haemog lobin inside red bl ood ce ll s.
fro m, individua l ce lls. Ma mma ls ue fa r mo re active tha n pla nts a nd
The mammalian circulatory By the end of this section you should be able to:
sy,;temcomistsofa
pump, many blood vessels a) state that the mamm alian circulatory syste m is a do ... d double circulation consisting of a heart,
andblood,whichisa bloodvessel, andblood
suspension of red blood b) obse rve and make pl a n diagra ms of th e structure of arte ries, ve in, a nd capillari'"' u,ing
cellsandwhitebloodcells pre pa redslidesandbea ble torecognise th ,ese v,esse lsusingthe lightmicroscope
in plamia. c) explain the re lationship betw~n th e st ructure and function of arte rii,s, V<!in• and capillarii,s
d) observe anddrawthe structure ofre dbloodce ll,,monocytes,neutroph ilsandlymphocytesu,ing
prepare dslidesandphotomicrographs
e ) state and explainthe diffe reoci,s betweeo blood,tissuefluidandlymph
fl describe t he rol e ofhae moglobinincarryingoxyge nandca rbondioxide wit hre ference tothe
role of carbonic anhydrue, th e formation of haemoglobink acid and carbaminohaemoglobin
g) describe a nd6plainthe significance ofthe oxygeodissociationcurvesofadultoxyhaemoglobin
atdiffe rentca rbond ioxide conce ntrations(the Bohr e ffect)
h) describe and6plainthe ,ignificance ofthe increaseinthe re dbloodce llcountofhuman, at
high altitude
151
El Transport in mammals
o pe n circulatio n of insects do uble circulati o n of m amma ls
bloodispumpedfO<Wards in the tuOOlarhea,~ aodtl>en bloodpas"-".ltwkethrough the
passesinthesin""-'S(openspac:es)between the0<ga,,s hearrinea<hcompletecirrulation
- ,.:.,,....,,..,-
~ ."'a"•"m r o
=>rnM•fy re:::::;,:::::::;""
'::.='lauooontJ>e · · · · __.. -
.. . i
two<:hamberedl>ean
Looking at these oiher systems helps us to understand the features of the mammalian circulation . h
becomes clear that the major advantages of the mammalian circulation are:
1 lnanopencirculation • simultaneous high pres.sure delivery of oxygenated blood to all regions of the body
therei1'littlecontrol • oxygenated blood reaches the respiring tissues. undiluted by deoxygenated blood.
over circulation'.
Suggest what this
means? Blood - th e transport medium
Blood is a special tis.sue of several different types of cell suspended in a liquid medium called
plasma(Figure8.2).
Plasma is a straw-coloured, very slightly alkaline liquid consisting mainly of water. It is the
medium for the continual exchange of substances by cells and tissues throughout the body.
Dissolved in the plasma are nutrient s in transit from the gut or liver to all the cells. Excretory
products are also transported in solution, mainly urea from the liver to the kidneys. So too are
hormones. from the ductles,; (endocrine glands), where they are formed and released. to the
tissues and organs
The plasma also contains dise;.ofrcd proteins. The principle blood prot"ein is albumin, which
has the role of regulating the water potential of the blood. The presence of dissolved a lbumin
stops too much water le"llv:ing at the capillaries by osmosis and also help, in the return of fluids
(Figure 8.8, page 159). Albumin and other proteins also assist in the transpon of lipids and iron in
the plasma. Other blood proteins present are antibodies (page 222) and components of the blood
2 Assumingthebody cloning mechanism.
containsSlitresof Suspended in the plasma are the blood = II~. lbe majority of these are n,cl blood cell~
blood,thatthereare (erythrocytes) - about 5 milHon in every cubic millimetre of blood. The red blood cells are formed
Smillion{S x 1o")red in the bone marrow tissue. Initially each red blood cell has a nucleus but early in development
bloodcellspermm'and thenudeusislostandthesetinycells(7µmindiameter)adopitheshapeofabiconcavedisc.
that a red blood cell Mitochondria and endoplasmic reticulum are also lOllt and a great de-al of a protein, hae moglobin ,
lastsfor120day1,how takes their place, together with the enzyme carbonk anhydras c . lbe role of mature red blood
many red blood cells
cells is exclusively the transport of respiratory gases in the blood. lbe lifespan of a working red
mmtbereplacedper
blood cell is only abou t 120 days, after which they are broken down and replaced. Most of their
day?
components are retrieved and reused.
152
8.1The circulate~ system I
f -~ o-e
redblocdcells w hita blood cells (leucocyt.. ) pl atelets
(e rythrocytes) (250000llr<n')
(smo11iontmm1)
I (lormootibodk".I) monocytes
phagocytes
»eutrophfs
(eng,u~baaefia)
153
El Transport in mammals
Table 8.1 Asumma,y of the roles ofbk,od components
Component
Trampo,tof
• nu tr ie nts fromthegutorl;...,,toallthecells
• excn,to,yp<odtKtssuc:has ureafromthel;,,e,tothekidoeys
• ho rmon esfromtfieendocrioeg!andstoall tissuesandorgam
• dissolved prote inswhochhaw,rolesindudingregulating theosmotKrnocent,abon
(waterpotentiaOoftheblood
• dissolvedpmteimtfia t area ntibodi es
Heat distributiontoal l tissues
Trampo,tof
(e rythrocytes) • oxyg e n fromthelur,gstonespifir,gcell,
• carbo ndioxidefmmresp;,ingcellsto tfielungs(alsocanied;nplasma)
Ly mphocytes; ma;o, roles in the immune syste m. indud;ng forming antibodies (Topic 7)
(le ucocytes) Pha gocytes:ingestbacteri aandcelllragments
Platelets Partofthe blood clottingmec ha ni sm
The hepatic
portal vein
The blood supply to the
liverisviathehepatic
anerybuttheliveralso
receives blood directly
from the small intestine,
vfaaveincalledthe
hepatic portal vd n .
Thisbringsmuchofthe
produasofdigestion,after
they have been absorbed
imothecapilbriesofthe
Figure8.3Thehumanbloodci,rulabon
vilHinthesmallintestine.
Notice that blood is pumped from the right side of the hean to the lung.~ . From the lungs, blood
retllfns to the left side of the hean. This pan of the system is called the pu lm o n ary dreula tlo n
From the left side of the hean, blood is pumped around the rest o f the body before returning to
the right side of the hean. This is called the ,.,·s tc m k d rculatio n
154
8.1The circulate~ system I
So you can see that blood passes tu.f re through the hean in every single circulation of the body.
This arrangement is a called a double circulation. This means there is no mixing of oxygenated
and deoxygenated blood in the hean, so the blood circulating around the body has a higher
concentration of oxygen than would otherwise be possible.
outerlaye,{tuoica
::1e:~1~i:fi~bresia)
/ / .. -.....:-.'\. of e!as:=Jt~~tlagen
muscle fibres
thehollowcl.'lltreof
atubeisthe lumen
'\.
'-::,,,...i;;;.,,;;,;;r;;;;
{inoer
;;:;-,1 ~~:;,~;..,""'"t
epithel;um - ,;ngle!ayer
ofcellswithsmooth;,me,
capillarywall{enlarged)
T5 uterya nd vein. LP(X20) - in section edmaterial(as herel. substanceslea..,(aod
,...,,........,..,,
veins aremore likely to appearsq uashed.whereas arte ries eote,)capillafies
uec"rcular·nserion thrnughthewal!s
Figure8.4Thestructun,ofthewallofarteries. ..,insa00capil!afies
15S
El Transport in mammals
Observing and drawing the structure of arteries.. veins and capillaries
You need to observe and draw the structure of aneries , veins and capillaries, using prepared slides
valve!sopenedby
and a light microscope under medium and high power magnifiaition, so that you will be able to
from behind
bloodJlfe<SUll'
recognise them.
Table8.20iffell'ncesbetweenarte,ies,cap;llafiesandvein,;
ir,dudingcOlltfactionsof
the muscles.which
=t
c=t
0\/u>rlayer(tunica extema) ofelasticfibres~ndrnltagen
Present - thin
!aye,
If you examine a prepared slide of an artery and vein under the high power of a microscope
you may see the layers of the walls in detail and you may be able to make out the muscle fibres.
Beaiuse of the low pressure in veins there is a possibility of back-flow here. Veins have ,·afr.,,. at
irnervalsthatpreventthis(Figure8.5).
We have noted that, as blood is transported around the lxxly the pressure of blood progressively
falls. Thisfactandtherolesoftheaneries.capillariesandveinsarereflectedinspecificchangesin
theirS1ructure(rable8.3).
Table8.3Thechangingstructureolbloodvesselsinre!ationtolunctkm
• Thewallsflaveahighprcportionofsmoothmusclefib,esandsoareabletoregu!ateblood
fi<1Nfmmthearteriesintothecapil!afies
Capillaries
Capilla ries: servethetissuesandcel!softhe • Narrow tubes. about the dia mete, of a sing~ !l'd blood cell (about 7µm). ,edoce the flow ,ate
body.Thebloodisunde,lowerpressull'(about toincreaseexchangebetweenbloodandtissue
35mmHg) • Thinwal!sofasingl,,!aye,ofendothelialcells
• Thewal!s havegapsbetweencellssufficienttoallowsomernmponentsofthebloodtoescape
andcOl\tributetotissueftuid(p,>ge146)
Ve ins: flave thin wall, ; the tunicaexte rna!sthe thkke,t !ayer.
Venules: collect blood ffom the tissues . They are • The walls cOl\sistof endothelium and a ve,y thin tunica media of a few scattered smooth
fo,medbytheu nkmolseve,alca pi lla,ies(p,essu,e musclefib,es
abou t 15mmHg).
Ve ins:receivebloodfromthe tissuesunderlow • Thetunkaexterna.coota iningelastkandrnl!.,.genfibres.!s present
pressure{p!l'e.sureabout5mmHg). • Thetooicameciamntainsafewelartic:fibresandmusde fibl!'s
• Theve in,;becomewider.solowefingpressure • Thep resenceofvalwspreventstheback-flowofblood
andincll'asingtheflow rate
156
8.1The circulate~ system I
Pressure: the pascal
(Pa)anditsmultiplethe
kilopar.cal(kPa) are 3 List the differences between the pulmonary circulation and the syi;temic circulation of blood
generally used by scientists 4 The graph in Figure 8.6 shows the changing blood velocity, blood pressure and cross-sectional
areaoftheve1,el1asblooclcirculatesinthebody. Usetheinformationinthegraphandin
to measure pressure
However, in medicine the Table 8.2 where appropriate to answer the questions below.
olderunit, millimetresof
mercury(mmHg) isstill
used(lmmHg = 0.13kPa)
Figure8.6Thechangw,gbloo<lvek>city.bloodpn,m,rear>dcros,-ll'Ct>ooalareaoftheve,selsasblood
cin:u!atesinthebody
a Comment on the velocity and pressure of the blood as it enter! the aorta compared with
whenitisabouttore-entertheheart
b What aspects of the 1tructure of the walls of arteries may be said to be in response to the
conditionofbloodflowthroughthem7
c What factors may cause the velocity of blood flow to be at the level it is in the capillaries
andwhyisthisadvantageous?
Table8.4Tr.msportrok>softhebloodcirrnlation
1S7
El Transport in mammals
Exchange in the tissues and the formation of tissue fluid
The blood circulatko delivers essemial resources (nutrients and oxygen. for example) to the tissues
of the body. lbis occurs as the blood flows through the capillaries, between the cells of the body
(Figure 8.7). There are tiny gaps in the capillary walls, found to vary in size in differen! pans of the
body. Through these gaps passes a watery liquid , very similar in composition to plasma. This is
li~suc fluld. However, red blood cells. platelets and most blood proteins are not presem in tissue
fluid. instead these are retained in the capillaries. Tissue fluid bathes the living cells of the body.
Nutrients are supplied from this fluid and carbon dioxide and waste products of metabolism are
carrieda,vaybyit.
tissuelluidf0<a>dimo
f;rnphcapillariesaod
~ !l!iesby
pre-capillarysphioo- - ' 1_ , surroondingti,sues
="o:~\'...c::; and by pressure
Th e roleofthe by.passve, .. 1
Whichtissuesolthebodyarebeiogservedby!hebkxxlcirrulatiooi,,e9uta1ed.Wheotioypre-capillarysphinctermusc:lesare
cootracted. blood now to that capillary bed is resuicted to a minimum. Most blood is then cfvened !hroucj, by-pass vessel, - ~ is
shumed1omhertissoesaod0<gar,sinthebody.
Fig u re8.7&changebem-eeobloodandcellsviatissueflukl
Given the quantity of dissolved solutes in the plasma (including and especially all the blood plasma
proteins), we would =pect the water potential of the blood to limit the loss of water by rumosis
In fact. we might expect water to be passing back into the capUlaries from the tissue fluid, due to
rumosis. However. the force applied to the blood by the heart creates enough hydrostatic pressure
to overcome rumOlic water uptake. at least at the aneriole end of the capillary bed. Here the blood
pressure is significantly higher than at the venule ends. Then, as the blood flows through the
capillarybedthereisprogressivelossofwaterandofhydrostaticpressure,too.Asaresult.muchof
the tissue fluid is able to eventually return to the plasma - about 90 per cent returns by this route
(Figure8.8).
So, funher along the capillary bed there is a net inflow of tissue fluid to the capUlary. However,
not all tissue fluid is returned to the blood circulation by this route - some enters the lymph
ca plllaries (Figure 8.9). Ly m1>h is the tissue fluid that drains into the lymphatic vessels, rather than
direcdyback into a blood capillary.
158
8.1The circulate~ system I
Bloodprotenl,p.articuiarfythealbumffi,c.mootl'SGil!"';theymaintainthewater"°tl'f11ialol
thepla,ma,""'ventingexcesslossofwateraodassi<lin,gtherenxnoffluidro !heG1pilla<ies
r
bioodin
-{- - -~-------
u•so~...........V--
ultraflltratlon dlffuslongradlent osmotic
of waterar.dsma ll (oxvgenarnlmmirots movement
molecules (oxygen, requi"l'dbyrespirin,g oiwate,
o,J
olwa\ef
!
dlffuslongradl • nt
(waste,,...tabolites)
hy drostatic
pressur<1
redocl'd
glucose,amioo cells)
acid<)dueto
hydrostatlc pressur•
_ /Inflow
Figure 8.8 ~on:esfor exchange in capil!afies
~
a lymphnode in section
~ ~
fa, J m
aa..,h•o•
engu!f ba<:teria,deb<is
- ~ \< - ~ :~:·:l~o~~::.,,,..m
(lymphocytesandplasmacelis)
are made here
K
the lympha tic circulation
theflow oflymph is propelledby
contractOOof,moothmuscle
fibresinthewalls ofthelymp hatics
andby them<>Yl'ment and
pre,.su...,ofthesurro,mdingtissue,
10 %.of Bac:kHowis preventedbyvalves
tissueAuid
bern=
inteslir,e(site
ofabso,pbon
ofprodurnof
lipodd>gesbon)
"""' :-,..-;'\_-~ ::,:c;::': del;ve,ed,resp irato,yga,es
··-
lymphatk
Figure8.9Thelayoutand roleolthelymphaticsystem
159
El Transport in mammals
Molecules too large to enter capillaries are removn:I from the tissues via the lymph system.
There are tiny valves in the walls of the lymph capillaries that permit this. The network of lymph
5 a What components capillaries drains into larger lymph ducts (known as lympha cks). Then. lymph is moved along
of the blood are not the lymphatics by contractions of smooth muscles in their walls and by compression from body
found in tissue fluid? movemetlts. Back-flow is prevented by valves, as it is in the veins.
b Constructatable Lymph finally drains 1:xick into the blood circulation in veins dose to the hean. At intervals along
to summarise the the way there are ly mph n odes present in the lymphatics. lymph passes through these nodes
differences between before it is returnn:I to the blood circulation. In the nodes phagocytk macrophages engulf 1:xicreria
tissue fluid and and any cdl detritus present. Also the nodes are a site where cenain cells of the immune system
lymph
are found. WerecurntothisissueinTopic 11.
Figure8.10Thestructu,eof
haemoglob;nariditsafftnitylor
160
8.1The circulate~ system I
A note on partial pressure Carbon monoxide poisoning
Weliveatthebottomofa·seaofair' - henceairpressure. Carbon monoxide is found in vehicle =haust gas, cigarette
lhe air we b=the is a mixture of gases and. in a g:,s mixture, s moke ,andasaproductofincompletecornbustionofnarural
eachgas=ensapressure.lnfact,thepressureofamixture gaswherethereisrestricredaccessforair.
of gases is the sum of the pressure of the component gases. Carbon monoxide combines irreversibly with the iron
Consequently,thepres.sureofaspecificgas ina mixture of of haemoglobin at the site that oxygen would otherwise
gases is called it5 pani.-..1 pressure. The symbol for partial occupy, forming earboxyhaemoglobin. The affin ity of
pressure isp and for the panial pressure of a gas X ispX carbon monoxide for haemoglobin is about 300 times
(so,forexample,,£0,denotesthepanialpressureofoxygen). greaterthanoxygen"sandatlowpanialpressuresofthis
lhe unit of pressure is the Pasc al (Pa) and its multiple the poison the blood"s ability to transport oxygen may be
kllopac<ca.l (kl'a) . fatally reduced .
Atmospheric pressure at sea level is approximately 100 (Incidentally,acatalyticconverter,whenfittedtoavehicle's
kPa and oxygen makes up about 21 per cent of the air. Wbat exhall.Sl.cansignificantlyreclucethequantityofthegasby
ispO,alsea/a'ei? oxidisingittocarbondioxide.)
Toretumtohaemoglobin.lJIOllthaemoglobinmoleculesare
fuUysaturatedatapan:ialpressuredoxygendonly8kPa
sooxygeniseffidentlyloadedundertherooditionsinour
ventilated lungs. However,inrespiringti:s.sue-thesiruation
isquitedifferem. Herethepaitialpres,uredoxygenis
muchlowerbecauseoxygenisusedin aerobicrespiration.
theoxyge,,dlssodalloo cllf'/e
olhaemoglobll\at lheCO, Oxyhaemoglobinwilldissocimeifthepanialpressureisless
concmtratlooilfOOOdrl'SJ"flng than8kPa,makingoxygenmoleculesreadilyavailableforuse
cffl,l.e.aboul9.31:PaCO, inthecellsdrespiringtissues
So0xygenisscarceinrespiringcdls.Onthe01her
hand.theconcentrationofcarbondioxideishigh. ln
thisenvironmentitisinterestingtoseewhateffectthe
concentration of carbon dioxide has on the loading and
unloading d oxygen by haemoglobin.
Jn fact. carbon dioxide has a marked effect. An
increased carbon dioxide concentration shifts the oxygen
dissociation curve to the right. That is, where the carbon
dioxideconcentrationishigh(o bviouslyintheactively
-\' respiringcdls).oxygenisreleasedfromoxyhaemoglobin
even more rC"lldily. This very useful outcome is known as
,,o,, the Bohr e ffect.
FigureS.11 Howcarbood>Oxide
faYOU"thereleaseofoxyger,in Carbon dioxide
respiringti,,..,... Carbon dioxide is uansponed in the blood. in lxxh the plasma and in red blood cells, mainly as
hydrogencarbonate ions
161
El Transport in mammals
a·,·n a lveolu s pl as ma
Oi inhighconcentfation 0., in ;r,coming air diffuses ;r,to red blood ce lls and combines
CO.,inlowrnoc:entration withhaemoglob;n{HH b)tofoonoxyhaemogk>b in(Hba:i)
, - - -.... Oi + HHba;=eeHbOi+ W
Oi ---- __ ) 1
tocamon dioxidegas
t issue fluid
Oiinlowrnoc:entration
~~:t:~
~----------------------
inthe,esp<ringtissoes
'
COi + H;,O ._.. HCOf + W
; _____ ___ bo_ · __ h,rdr __ , --- CO.,
162
8.l lhehean .
Theresultofthesechangesisthatasaltitudeincreasesitbecomesincrea.singlyhardfor
haemoglobin in red cells in the lungs to load oxygen. Once the percenmge saturation with oxygen
is lowered, this is detected by chemorecepcors. The response of the respiratory centre is to cause
the taking of extra deep breathes. As a resuh, more carbon dioxide is lost from the lxxly, which
causes a small but significant rise in the pH of the blood. Now the chemoreccpiors become
ineffective. Ventilation regulation is hampered.
The body cannot adapt to high altitude immediately; sudden, prolonged exposure at very
high altitudes by people without prior =perience of these conditions can be fatal. However,
progressivdythefollowingehangestakeplacc:
• A more alkaline urine is secreted by the kidney tubules via the collecting ducrs, and the pH of
the blood returns to normal. As a result, the carbon dioxide chemoreccpiors become sensitive
again, and normal ventibtion is maintained.
• Bone marrow tissue, the site of red cell formation, produces and releases more red cells, thereby
enhancing the oxygen-earrying capacity of the blood (fable 8.6).
Table8.6Thechai,.gei11redMoodcellcoontinhumamatdifferentaltitudes
O(sealevel)
500()• asatempo,ary,,;.;rtor
These changes are called 'acclimatisation '. Other mammal species have evolved at high altitudes.
such as the llama of the mountains d South America. In this case the mammal has a form of
haemoglobin that loads more readily at lower partial pressures d oxygen.
The mammalian heart is By the end of this section you should be ab le to:
adoublepump:theright
side pumps blood at low a) describe the exte rn al and inte rnal structure of the mammalian h..art
pressuretothelungsand b) explainthe diffe rences inthe thicknessof the w a llsofthe diffe re ntchambe rsinte rmsoftheir
theleftsidepumpsblood functionswithre fe re nce toresistance toflow
athighpre11uretotherest c) de scribethecardiaccyde(includingbloodpre"urechangesduring~oleanddiastole)
of the body. d) explainhowheartactionisinitiatedandcontrolled
163
El Transport in mammals
contract to all cells simulrnneously. The impulse to contraa is generated within the heart muscle
itself (known as a myogc nic o rigin). nOI by nervous stimulation (a n c u cogc nic origin ). Hean
muscle fibres contract rhythmically from their formation until they die.
Figure8.13Ca,di.Jcmuscle
164
8.llhehean .
- - ~ " l -- - - -1enr,umona,y
....,
Figure8.14The,tructureoftheheart
165
El Transport in mammals
HeartvaM.'<aloNbloodllowin Atria l systole
ooedirectiooooly.They""' • A!Jialwaliscootract - !hep<eSSUJeisraisedintheatria
opened and dosed by • \ol>nara,aeandpulmooa,y,,..insa!l'dosed
diflefl'J>Ces inbloodp,essure • At1io,,..mriru~valvespusl>edopro - bloodllowsintothe,,..mrides
• They,.,.,openedwheritr>e • \ol>ntriculMwallsre!axed - thepressu!l'islowinthe,,..mrides
Pf'-"'lleOO!hejnpulside •CHghpressureintheaortaandp;,jmooaryaneries(duetotr>eelaslic
(causedbyooosclecoouaaK>Jl) andrrusc:lelibresintheirwal s) - thesemiUnarvalves""'shut
exceeds!hatootheootpmside
• They..,,.dosedwheolhe V""triaJlar systole
pressureootheootputside " :':'n~:~alisrort!Jactstrongly theplt'SSUfe1Sra,sed n
exceedsthatoothe;nputSOde
~=~·o~u~~the
input side
-~=":.e~. ~~;,:,~~:;,r::.!:°'
ll'laxed - thep,essureinbo!hatriaar>dvent1ic:iesisk,,v
Figure8. 1!tTheitcbonoftheheartvalves
166
8.llhehean .
lbe atrium then relaxes for the remainder of the cycle (atrial diastole , about 0.7 seconds).
Next the ventricle contracts (veniricular systo le. about 0.3 seconds). The high pressure this
generates slams shut the atrioventricular valve and opens the semilunar valves, forcing blood
into the aorm. A "pulse', detectable in aneries all over the body, is generated (see below).
This is followed by relaxation of the ventricles (vcmricuiar Jias1olc, about 0.5 seo:inds) until
Each contraction of cardiac muscle is followed by relaxation and elasiic recoil. lbe changing
pressure of the blood in the atria, vemrides, pulmonary an~ and aorta (shown in the graph in
Figure8.16)automaticallyopensanddruesthevalves.
-
musd escootract,
pushing blood pas!
musd .. rom,aa,
causongbloodp<e:<sure
todose!hebicuspid
valve and open the
<emolunarvalves,
-
ventridaand
atriummuKles
relax,whilett>e
:~o~~:::
~~~,~~. . ~=.~~
atrial systo le
Figure8.16Tfiecardiaccyde
167
El Transport in mammals
Heart rate and the pulse
The contmaion of the ventricle walls forces a surge of blood into the aorta and the pulmonary
8 Examinethedataon arteries come under great presslU"e. This volume of blood is called the s troke ,·olun1e . Each surge
pressure change during stretches the elastic fibres in the aitery walls (fable 8.3, page 156). The anery waUs are distended
therndiaccydein
as the surge passes, before the subsequent elastic recoil occlU"s. This is known as the pulse. Each
Figure8.1S.De1etmine
contraction of the ventricles gene-rates a pulse, so when we meaSW"e our pulse rate we are measuring
orsuggestwhy
a pressureintheaorta our heait rare. We can measure heait rate in the caro1:id artery in the neck or at the wrist, where an
isalways§ignificantly artery passes over a bone. Incidentally, the amount of blood flowing from the he:iit is known as the
higher than that in rnrdfac o utpm . At rest, our cardiac output is typically about S litres of blood per minute.
the atria
cardiac output = stroke volume X pulse rate
b pressurefall1
=tabruptlyin Pulsation of the blood flow has entirely disappeared by the time it reaches the capillaries. This is
the atrium once due to the extensive natlU"e of the capUlary networks and to the resistance the blood experiences as
ventricularsystolei1 itflowsthroughthecapillarynetworks.
underway
c thesemiunarvalve
intheaortadoesnot
open immediately
Origin and control of the heart beat
The he:in be-:its rhythmically throughout life, without rest. apan from the momentary rebxation that
that ventricular
occurs between each beat. Evenmoreremarkably,hea1tbeatsoccurnarurally,withoutthecardiac
systole commences
muscleneedingrobestimularedbyanexternalnerve. Sincetheoriginofe:ichbeatiswithinthe
d whenventricular
hean itself. we say that he:in be-:it is 'm yogenk" in origin. However. the alternating contractions
diastole commences,
of the cardiac muscle of the atria and ventricles are o:introlled and co-ordinated precisely. Only in
there is a significant
delay before the this way can the heait aa as an efficient pump. The positions of the struaures within the hean that
bicuspid valve bringthisaboutareshowninFigure8.17.
opem,despite The steps in the control of the cardiac cycle are as follows.
nsmgpressurem
• The hean be:it originates in a tiny pan of the muscle of the wall of the right atrium, called the
the atrium
s inoatrial node ( SA node) or p acemaker.
e itissignificantthat • From here a wave of excitation (electrical impulses) spreads out across both atria .
about SO percent • In response, the muscle of both atrial walls contraas simultaneously (m rial systole ).
of the cardiac cycle
• This stimulus does nOI spread to the ventricles immediately because of the presence of a narrow
is given over to
diastole band of non-eonducting fibres at the base of the atria. These block the excitation wave, prevent-
ing its conduction across to the ventricles. Jrutead. thestimulusispickedupbythea trlo \"e n -
trlcular node (A\' node), situated at the base of the right atrium.
• After a delayof0.1---0.2 seconds, the excitation is passed fromtheatrioventricular
stimulation of heartbeat
node to the base of both ventricles by tiny bundles of conducting fibres known as
-
originates inthe musda
the Purkyne tissu e . These are collectively called the bnndks of Hi~
sinoatrial • On receiving this stimulation from the bundles of His, the ventricle muscles stalt to
contract from the base of the heait upwards (, ·entrknlar systok).
(pacemaker}
;,,rqn • The delay that occurs before the atrioventrk,.1lar node aas as a 'relay station' for the
impulse permits the emptying af the atria into the ventricles to go to completion and
preventstheatriaandventridesfromcontractingsimultaneously.
• After every o:intraaion, cardiac muscle has a period of insensitivity to stimulation,
the re fractory pe riod (a period of enforced non--contraaion - dias tole). In this
phase, the he:in begins to refill with blood passively. This period is a relatively long
oneinhe:11tmuscleandenablesthehea1ttobeatthroughoutlife.
168
8.llhehean .
Electrocardiography
The impulses (aaion potentials) that originate in the sinoatrial node (pacemaker) of the hean
duringthecardiaccydeproduceelectricalcurrentllthatarealsoconductedthroughtheOuids
of the body as a whole and can be detected at the body surface by electrocardiography. Here,
electrodesareanachedtothepatient'schestandtheelectrical activitydetectedisdisplayeda.s
an electrocardiogram (ECG) by me-ans of a chart l'<."C<X'der (Figure 8.18).
Electrocardiography has clinical applications: it is an aid in the diagnosis of cardio,...,.cular
disease (CVD, page 198). Some heart conditions that can be detected by the analysis of
electrornrdiogramsarelistedinTable8.7.
normal • l•ctro<ardiogram(ECG).analysed
- = ,.,
- atriajromraaioo(atrial systole)
~ ""mriru~romraaion(sys\ok,)
::~=~Dll-leamm
timef0<;~se!ObeID11ducted
::"~oodemventrides,via
QRScomp lu ::,:1i=!';~t;c~=tioo-
Vl'<1lriru~rl'polar;.;.,1Klfl -
fl'laxalioophase
Q-T mterval
1 ta<hytardia ~
heart rate,Sove<
100 beatsfmonute
2ventrku larfibrillation ~
uncontrolledromractioo
oithe,,,..mrides - lrttle
bloodisp<.<nped
(Contim,ed)
169
El Transport in mammals
(Cominucd)
Table8.7Hearlc<><>ditionsdetectedbyECGanal)'Sis
Heartconditionsdetectedbyabnormal ECGtraces
Tach)'(a,rlia A normal .tdult heart beats between 60 and 100 times a minute; a heart rate ove, 100
beatsam;nuteisca lledtach)'(ardla
Tach)'(ardla may be ,e!atively harmk>ss and need oo ITT'atment but some forms can be
life-threatening
Asynchmnousrnntracbonoftheventric:lemusclefib,esresultsinafailureoltheheart
to pump suffic.,nt blood because some muscle fibres are contracting whilst othe" are
relaxing
Themostrnmmoositeofbkxkageisattheat,ioYl'l\tricu!a, node
Anhythmia Arrhythmiaisarnnd~ionofirregularrtyinheart,hythmdueto a defectinthe
rnnductionsystemoltheheart. ltmaybedueto
• drugs.suchasnicotineo,akohol
• anx.,ty.hwathyroidismo,potassiumdeficiency.
170
Examination style questions I
Examination style questions
1 Fig. 1.1 shows the changes in blood pressure in the left 2 a) What tissue is found in all types of blood vessel? How it is
atrium,leftventrideandaortaduringonecomplete adapted to its functions? [4]
contri!dionoftheheart.ltalsoshowsarecordingofthe b) Ina table, give the differences between the structure of
electricalactivityoftheheart anarteiyandaveinasseenintransversesection [4]
c) Explain the difference betweenasingleandadouble
circulation,andthebenefitsgainedbyrespiringlil1ues
Key: served by the latter. [2]
d) lnatable,givethedifferencesincompo5.itionbetween
plasma and tissue fluid [4]
e) Listinorderthepathwayofvesselsandchamber1taken
byaredbloodcell,fromleavingtheliveruntilitreaches
cellsofthemusclesofthearm [6]
[Total: 20]
Fig.1.1
a) Namethesourc:eoftheelectricalactivityintheheart. [1]
b) Explainhowtheheartisco-ordinatedsothattheventricle
contractsaftertheatriumhascontri!ded [4]
c)WithreferencetoFig.1.1,calculatetheheartratein
beato;perminute. Showyourworkingandexpres1your
answer to the nearest whole number. [2]
d)Thepre5sureintherightventriclei1rarelyhigherthan
4.0kPa.
Explainwhythepressureintherightventricleismuch
lowerthanthatintheleftventride [2)
[Total: 9]
(Cambridge International AS and A level Biology 9700,
Paper 02 Q4 November 2007)
171
AS Level
172
9.1 The gas exchange system I
outer surfaces are no longer suitable for gas exdumge and the organism requires an alternative
res piratory s urface. Active organisms have an increased metabolic rate. too, and the demand for
1 listthreefactor.; oxygen in their cells is higher than in slow-moving or irutctive organisms. So we find that, for many
affecting the rate reasons,largeractiveanimalshavespecialisedorgansforgasexchange.
of diffusion across a Efficient respiratory surfaces in animals take various forms, such as thegUlsoffish, the lungs
~urfaceandexplainwhy of mammals. the tubular system of many insects that carries air to the most acrively respiring
eac:hiss.ignificant.(You organs.Allthesesystemsprovidealargethinsurfacearea,suitableforgasexchange. In addition,
mayneedtoreferto conditionsfordiffusionareoftenimprovedbythreerefinements
Table4.1,page79.)
l A \"cntll:uion m<-'Ch:1nism - a pumping mechanism that moves the respiratory medium (water
2 Ouetothepresenceof
or air) over the gills or into and out of the lungs or tubes. This maintains the concentration
haemoglobin, about
20cm1 ofoxygenis gradient for diffusion.
carriedper100cm1 of 2 A blood circulation syste m - a means of speeding up the transpon of dissolved oxygen from the
blood.BycOfltrast,the respiratory surface as soon as itha.s diffused in. This,too, maintains the concentration gradient.
solubility of oxygen in 3 A ha,, m protein, such as haemoglobin. that can associate with oxygen and so increase the gas-
waterisonly0.02Scm' carrying ability of the blood. For example. we have seen that our blood contains ttd blood cells
percm'underthesame packed with the n,splratory pigme nt haemoglobin (Figure 8.10, page 16o).
cOflditions. HowmlKh
more oxygen is carried The lungs of mammals
byalitre{lOOOcm')of
We can now consider the human lungs as strucrures adapted for rapid gas exchange. The structure
blood compared with
of the human thorax is shown in Figure 9.2
the same quantity of
water? The lungs are housed in the thorax , an air-tight dome-shaped chamber formed by the rib cage
and its muscles (l nt cn:ost:tl muscks) and with a domed floor, the diaphragm. lbe diaphragm
is a sheet of muscle attached ro the body wall at the base of the rib cage, separating the thorax
from the abdomen. lbe internal surfaces of the thorax are lined by the pleurnl membrane,
which secretes and maintains pleural fluid. Pleural fluid is a lubricating liquid made from the blood
plasma. This fluid provides the surface tension that holds the lungs ro the rib cage and pro1:eas the
lungsfromfrictionduringbreathingmovements
nas,1lc.,,.;1y
palate
bucalcavity
epiglott;,; '"'"'"'
'"'"'
~\IS
pleural membranes
intercostal
muse~ pleural cavity
ointerllill ~';' tainin 9plemal
.exterllill 0
lung
di.lJ)h,agm ::i~ t
c"'ity
Figure9.2Thestructu,eofthehum,mtho,ax
The lungs connecr with the pharynx at the rear of the mouth by the trachea. Air reaches the
tracheafromthemouthandnostrils,passingthroughthelarynx('voicebox").Entryintothelarynx
is via a slit-like opening, the glottis. Above it is a canilaginous flap, the epiglottis. Glottis and
epiglottisworktopreventtheentryoff<:xxlintothetrachea.
173
IJI Gaseousexchangeandsmoking
The trachea then divides into two primary bro n c hi, one to each lung, Within the lungs the
primary bronchi divide into secondary bronchi and these continue to divide into smaller bronchi
a branching system referred to as the 'bronchial tree', lbe smallest bronchi themselves divide into
bronchiolc,i, The finest bronchioles, the terminal bronchioles, end in air sacs - the aln,oll,
SEMshowingthattheliningofthebronchiolesisofsimple,
ciliatedcolumnarepithelium.withm11<ussecretedbygobletcells
(x6000)
,.,
,;mple.columnarepithelium
goblet cel l
Ooeofthemanywaystobacrn
smokedamageslungtissueis
due to its temporarily
anaesthetisingeffectoodlia
movements. MtKusloadedwith
harmfulpollutantsacrnmulates
ontheepitheliumcells.causing
partialblodingoftheairways
Figure9.3Cjliatedepitheliumwithgobletcell,
The trachea lies beside and in front of the oesophagus (the 'food tube"). Any hard mass of food
passing down the oesophagus might interrupt the air supply to the lungs. Incomplete rings of
cartilage in the trachea wall prevent collapse under pressure frcm a large bolus passing down the
oesophagus(Figure9.4).
174
9.1 The gas exchange system I
The walls of the bronchi and larger bronchioles, in addition to rings or tiny plates of rnnilage,
also contain s mooth mus.de . At each division of the bronchi3J tree the amo,mt of canilage
decrea~ and the amount of smoo1h muscle increases. Together they prevent collapse of these
tubes - collapse might be triggered by the sudden reduction in pressure that occurs with powerful
inspirations of air, for example. The smood, muscles also regulate the size of the smaller airways as
The walls of the smaller bronchioles are without canilage (Figure 9.5). lbese tubes, the
narrowest of the airways serving the alveoli, branch repeatedly and, as they do so, become
progressivdy narrower. Here, smooth muscle is the major component of their walls and the lining
of columnar epithelium conrnins only very occasional goblet cells.
So the air sacs are supplied with warm moist clean air for gaseous exchange, but of course the
lungs cannoc prevent some water Joss during breathing a significant issue for IJIOllt terrestrial
organisms(Figure9.6).
photomicrographoftrachea inTS(,c10)
lin;ngofciliate
epithelium with
goblet cell,
smoothmusclejoink,g
cartilage end,
cilia+columna,epithe!ium
w ith goblet cell,
partofC-,haped
cafti\agering
Figure9.4Thestructu,eofthetrachea
175
D Gaseousexchangeandsmoking
176
9.1 The gas exchange system I
Bronchiole structure and asthma
Asi hma is a disease of the airways of the lungs. Jn an as1hma attack, the bronchioles are narrowed by
excessive contracrion of the smooth muscle in their walls. Immediately, getting air into and OIJI of the lungs
becomes difficult. Also, extra mucus is produced, exaggerating the symptoms. Breathing can become very
difficult indeed.
Aslhrna attacks may be staned by the arrival of irritants like pollen, dust from pets or droppings from
house dust mites. Certain viruses a nd the oxides of nitrogen present in vehicle exhaust fumes may also be
triggers
During times o f physical aaivity our involuntary nervous syscem naturally releases hormones thm relax
the smooth muscles of the bronchioles, dilating them so that air reaches the alveoli more quickly. Medical
treaunents for asthma sufferers seek to trigger this relaxation, too (Figure 9.7).
~n=~\efCOS!al
musde<cootract - fib<
::~::~~\ef((lS~
musclelre!ax - nb<
""""'upwardsar<I moYe-.W..-dsand
ootwa rds. andlhe Inwards.and the
dlaphragmclown dlaphr,q,,up
•::;:m ·!1:1~ag,:.,,
~·-
(rlb<artlrulate
'M!hlhe!ho<ack
vefte,bfae)
oo>
pleual!Md
<teroom(ma<t
rtisareatt.lched
lungtl,sue herebycarll~)
d"'!)!l,agm
~meo!lhettiorax(ar>dthefefoo,oflhe voklmeoflhethofax(andlhere!oreo!lhe
lu"9s)l.-.:reases;11n,,1mel,reducedbek>N lung1)<lecreases;J)fes:sure~lncreasedilOOW'
atmosplwrkPfessurear<lal<flowsln atmolp!>eflcJIJes1Y,ear>dal<!b,v,;oot
Figure9.8V..nti!ahooolthelungsinquietbreath;ng
177
IJI Gaseousexchangeandsmoking
Figure9.9Theroleofelasticlibresinthea~eoliandbronchioles
A capillary system wraps around the dusters of alveoli. Each capillary is connected to a branch
of the pulmonary ar1cry 3nd is drained by 3 branch of the pulmonary vein (Figure 9.10). The
pulmonary circulation is supplied with dcoxygcnmcd blood from the right side of the heart and
oxygenated blood is returned to the left side of the heart to be pumped to the rest of the body
(page165).
There are some 700 milHon alveoli present in our lungs. providing a surface area of about 70 m'
in total. This is an area 30--40 times greater than that of the body's external skin. The wall of an
alveolus is formed by pavement epithelium, one cell thick. An epithelium is a sheet of cells bound
strongly together, covering internal or external surfaces of multicellular organisms. Lying very dose
is a rnpillary, its wall also just a single, Oattened (endothelium) cell thick. The combined thickness
of walls separating air and blood is typically 2--4 pm thick . The rnpUlaries are =tremely narrow,
just wide enough for red blood cells to squeeze through, so red blood cells are dose to or in
contactwiththecapillarywalls
Theextremelydelicatestructureofthe alveoli is protected bytwotypesofcell. present in
abuodanceinthesurfacefilmofmoisrure(Figure9.IO).
• Macrophages (dust cells) are the main detritus-collecting cells of the lxxly. lbey originate from
bone marrow stem cells and are then dispersed about the body in the blood circulation. The...,
amoeboid cells migrate into the alveoli from the capillaries. Here these phagocytic white blood
cells ingest any debris, fine dust particles, bacteria and fungal spores present. They also occur
lining the surfaces of the airwaysleadingtothealvcoli.
• Surfactant cdls produce a detergent-like mixture of lipoproteins and phospholipid-rich secre-
tion that lines the inner surl'ace of the alveoli. This lung surl'actant lowers surface tension, pennit-
ting the alveoH to Oex ea.sily a.s the pressure of the thornx falls and rises . ltreducesatendeocyof
alveolitocollapseonexpiration
178
9.1 The gas exchange sys1em I
photomicrographofTS • lvtoll,HP
____ ,...,,
., ..
:. -r :;-:.:~_·~ sr:~,
7
position of
elutic lim,,
IJ 1 1
illwiol;orw;ol
• \
Cilpillilry
(S11u;omous epithelium)
Flgu .. t.10Gasexchangeond'Ol!ilYe~;
:tbbod
179
IJI Gaseousexchangeandsmoking
Blood arriving in the lungs is low in oxygen (it has a lower partia l pressure of oxygen than
thealveo brair - seeTable9.l)buthighincarlxmdioxide(ithas3higherpartfalpressureof
6 Explainhowthe carbon dioxide th3n the alveolar air). As blood flows past the alveoli, gaseous =change occurs by
ITldCrophagesremove diffusiOfl. Oxygen dissolves in the surface film of water, diffuses across into the blood plasma and
foreign matter around into the red blood cells where it combines with haemoglobin to form oxyhaemoglobin. At the same
them.{Thedisc:us!>ion time, carbon dioxide diffuses from the blood into the alveolus.
onpage222in Air flow in the lungs of mammals is tidal in that air enters and leaves by the same route.
Topic11 mayhelpin Consequently there is a resid ual volume of air that cannOI: be =pelled. locoming air mi=s with
the answering of this anddilutestheresidualair. The effect of this isthatairinthealveolicontainssignificandyless
question.)
oxygen than the 3tmosphere outside (Table 9. 1). Nevertheless, the lungs are efficient CO"gans of
7 Createatablelisting
gaseousexch3nge.
the main structural
features of the air
saa{alveoli}and Table9 .1 Thernmpositicmola ir intf>elur,g,
identifying the effects
and consequence of lln spiredair I Alveolar air I Expiredair
each feature for gas o~,., 1 20% 1 14% I 16%
exchange . I I s.s%
8 lfthemncentrationof
o.04% I 4.0%
carbondioxide~reto
Nitl'Ogen 1 79% I 81% 1 79%
buildupinthebloodof Wate,vapour I variable I saturated I saturated
a mammal, why would We flaw, alre;,dy noted Iha~ in a mixture of gases, ea<l, component gas exerts a panial pressure in prcpoftion to
thisbeharmful7 howmuc:hispresentTf>epartialp,essureofoxygeniswrittenaspO,andofca,bondioxideaspCO,
Atsealevel.theatmosphericpressureis101.3kPaso.fOfexample.thepartialpressureofoxygenintheairat
sea level is
pO, -
1~~3 x20 - 20.3kPa
butinthealveolusthepartialpiessureofoxygenisonly
pO, - l~~J X 14 - 14.2kPa
180
9.2Smoking .
9.2 Smoking
• C.-trdn ogcn s In the "tar" com ponent , of which there are at least twenty differ=t types.
Panicularly harmful are cenain polycyclic aromatic hydrocarbons and nitrosamines. Remember.
a carcinogen is any agent that may cause cancer by damage to the DNA molecules d
chromosomes. Such 'mistakes" or mutations d different types may build up in the DNA of body
cells=posedtothescsuhstances
• Nicoti n e. a stimulating and relaxing drug which, on entering the blood stream, is able to
cross the blood-brain barrier. In the brain it triggers the release of dopamine, the natural
neurorransminersubstance(page319)associatedwithourexperienceofpleasure. Long-term
=posure to nicorine eventually comes to have the reverse effect, actually depressing the
ability to experience pleasure. So more nicotine is needed to 'satisfy', and cigarenes become
addictive; it is as addictive as heroin and cocaine, in fact. Smokers find it increasingly hard to
q uittheh3bit .
Nicotine 3Jso increases the hean rate and blood pressure. It decre3ses the blood llow,
panicularly in the hands and f=t, and it makes blood clotting more likely. 1t can make platelets
stick together.
• C:u-bon mon oxid e , a gas that diffuses into the red blood cells and combines irreversibly with
haemoglobin (page t6o). Jn smokers, the blood is able to transport Jess oxygen. The strain this
putsontheheanandcirculationismostappareruduringphysicalactivity.
Carbon monoxide also prcmores the release of damaging free radicals and makes platelets and
ncutrophilssticktogether.
181
IJI Gaseousexchangeandsmoking
Cigarette smoke reaches the smoker's lungs when it is drawn down the cigarette and inhaled, bm
it reaches ocher people, {O(), when it escapes from the glowing tip into the surrounding air. These
latter fumes normally have a higher concentration of the toxic ingrediems and it is this mixture that
ochers inhale. 'Passive s moking' has itself been shown to be dangerous, 10().
Two of the major diseases that are directly induced by cigarette smoke are c hro nic oboi tru c tin,
pulmo na ry disease (COIID) and lung can c er. We will review each in tum, focusing on the
effects of tar and carcinogens in tobacco smoke as the rnusative agents.
Emphysem a is a disease in which the walls of the alveoli lose their elasticity. It results in the
destruction of the Jung tissue with time. This is bemuse such lungs now contain large numbers of
macrophages, accumulated from the blood circulation (Figure 9.10). These phagocytic cells release
a high level of the natural hydrolytic enzymes that break down elastic fibres of the alveolar walls
With failing elastic fibres, the air sacs are left over-inflated when air becomes trapped in them and
they fail to recoil and expire air properly. Small holes also dt,>velop in the walls of the alveoli. These
begin to merge, forming huge air spaces with drastically lowered surface area for gas exchange
The patient becomes permanemly breathless. The destruction of air sacs can be halted by stopping
smokingbutanydamagedonetothe lungscannocbereversed.
Figure 9.12 SEMs of Healthy human lung ti=e Oeft) and human lung tis,i,e showing advanced emphysema (right)
182
9.2Smoking .
Lung cancer
Persistent exposure of the bronchi to cigarette smoke results in damage to the goblet cells .
These are then repbced by an abnormally thickened epithelium. With prolOllged =posure to the
carcinogens, permanent mutations may be triggered in the DNA of some of the cells Ulat have
replacedgobleccells. lfthisoccurs intheirprom--oncogenesorrumour-suppressinggenestheresult
islossofcontrolovernormalcellgrowth(page 104).
A single mutation is unlikely to be responsible for triggering lung cancer, it is the accumubtion
of mutations with time that causes a group of cells ro divide by mitosis repeatedly. with01Jt COfltrol
or regubti=, forming an irregular mas., of cells - the mmour. Tumour cells then emit signals
promOl:ing the development of new blood vessels to deliver oxygen and nutrients, all at the
expense of the surrounding healthy tissues. Sometimes tumour cells break away and are carried to
cxher pans d the body, forming a secondary rumour (a process called mct.-.s1a.sis). Unchecked
cancerouscellsultimatelytakeoverthelxxly,leadingtomalfunctionanddeath.
183
IJI Gaseousexchangeandsmoking
,,,~
~ ~.~ .
0.4
patient.onwhattypeoflungcancerhasbeencontracted. andhowfarithas
l 0.2 • --P
spread. Many of the rumours detected by early diagnosis may be removed by
surgery or suppressed by radiotherapy (with X-rays or another form of radiation) -
possibly followed in either case by chemotherapy using anti-cancer drugs.
0
o 5 10 15 20 25 However, morethanhalf ofpatientsdiagnosedcurrentlydiewithinayearbecause
yearsafte,gMngupsmoking lungcancerisadeadlydiseasethatisrarelydiagnosedearlyenough. Incidentally.
figure9 .14Deathrateslromc.:,r,ce,afte,givingup apersonwhogivesupsmokingmayeventuallyachievealife=pectancyvery
smoking similartothatofanon-smoker(Figure9.14).
Table 9.2 Longevity in smoke,,; ar,d r,oo-smokefs Table 9 .3 Car,ce, rates and the numbe, of cigall'ttes smoked pe, day
... Nurnberofpeoplestillaliveateachage
Light m,okers Hewy smokers
Numberofcigarirttes
...-.okedpe,d.ly
lncidenceofcancer
per100000men
A survey of smoking in the UK was commenced in 1948. at which time 82 percem dthe male
population smoked and d whom 65 per cent smoked cigarettes. This had fallen to 55 per cent by
1970 and continued to decrease. In same period the numbers d females who smoked remained
just above 40 per cent umil 1970, after which numbers also declined (Figure 9.15). Look carefully at
thechangingpatternintheincidencesoflungcancer - figuresareavailablesince1975
Concerning smoking and n,splratory JlseasC'!l, the National Institute for Health and dinical
Excellence (NICE) reports that chronic obstructive pulmonary disease (COPD) accounts for
30 OOO deaths per year in the UK, of which 85 per cent could be attributed to smoking - a figure
approximately double the European average. The World Health Organisation (WHO) provides
leadership on health maners for the United Nations by monitoring and reporting on the health
of peoples and assessing health trends worldwide. lbe World Health Organisation predicts that
by 2030, chronic obstructive pulmonary d isease will be the third mOllt common cause of death
worldwide
184
9.2Smoking .
- matesmokingp<evalence - femalesmokingprevalence
matelungcancerincklence female lung cancer incidence
9 LookatFigure9.1S
Comment on the
incidence of lung cancer
in men and women
between1975and
2007 in relation to
the changing pattern
of smoking since UK
records began
185
IJI Gaseousexchangeandsmoking
The impacts of this effect on the vascular system are raised blood pressure and a reduced oxygen
supply to the major organs and limbs . At the same time, the enhanced release of the hormone
adrenalin causes an increased heart rate. which in these circumstances, contributes to raised
blood pressure, too. A condition of permanently raised blood pressure is known as hypertension
(sre 'Blood pressure and its measurement' below). The epidemiological evidence is of an
increased risk of coronary artery disease at an earlier age in smokers than in non-smokers. In fact
the first evidence of the link between smoking and cardiovascular disease (CVD) established this
relationship.
It was in 1950 that Dr Richard Doll and colleagues. working at St Thomas"s Hospital, London,
publishedtheresultsoftheirinvestigationofthecausesofdeathduetocardiovasculardiseaseina
large &le of working doctors (am011g whom the habit of smoking was widespread at that time).
These implicated smoking as the culprit (Table 9.4). Today, few doctors smoke.
Table9.4Mortal1tyfromcardffiasrnla,diseaseamor,gasampleofdocto,s
Causeofdeathinasampleof
Cigarette smokers
"'""'°"""
COfOflaryheartdisease
AneU')"ID
186
9.2Smoking .
High-nmmal
Stage1hypertension
f = - c - ' - - - - + ~ ~ - - - +"~ ode
= ''"
~ -= ~ ') ~h""
= '=~·'----, Treatmentessentialilthesecorn:litionsper,;ist
Severe(stage3)hwertension
Ver"jsevere(stage4)hypertension
L>{ .\J ~
(kPa)aregenerally
usedbyscienliststo
inmedi::.':, the ~
OO~ofp<l'SSUfl'.
"m ilimetreolmm:Uly'
M=>>•"'"
(mm l-lg)isstil used
(1mml-1g=O.\JkPa) """'
p<essure
hand pump
187
IJI Gaseousexchangeandsmoking
Smokingisdearlyahealthriskwithlife-threatening
consequences. Perhaps the best response is to pre-
ventthe problemsarising bypersuasion, OOckedup
bylegislationwherethisisthoughteffective.One
developedcountry"s current response to the issue is
torestricrtheadvertisinganddisplayoftoOOcco
products. enforce bold , unavoidable health warnings
on cigarette packets and outlaw smoking in public
buildings and workplaces. At the same time, the
authority responsible for public health is assisting
thoseaddictedtonicotinethroughsmokingto dis-
cardthe habit if they can. This sustainable 'carrot and
stick' approach is having some significant positive
outcomes but there remain many who continue to
smoke.
Meanwhile, throughout the world there continues
Figu ,.., 9.18 Pmuasiw,advertis.ing.itseemsl tobehugeinvesanentsbythetobaccoindustry
in the growing of the crop, the manufacturing of
'attracrive"packagingfortobaccoproductsandthe
placingofadvertisementsincountriesandplacesthat
are open to them .
11 Cardiovascular disease
wasnotanissuefor
people before the
twentieth century.
Todayitisamajor
health issue.Make a '"'""
~
conci1e,annotated ~ 1500
listofallthefactor1 8
thatyouthinkmaylie 1220 1 2'00'
behind this change
touseinagroup
diKussionwithyour
"''"
~"
Fig un, 9.19 Worldtrends . Hl,j>lncomecOUf\trles . l.owandmld<telncomecOUf\trles(exdudlngChlM)
in per capita cigarette
consumpt·on
188
9.2Smoking .
Whilst the number of people smoking cigarettes is actually faUing in developed countries, smoking
is Ofl the rise in the developing world (Figure 9.19). The global heahh issues generated by cigarette
smoking are d epidemic proportions - about one-third of the global adult male population smokes.
Among the World Health Organisation a""3.'l, the EaS1 Asia and the Pacific Region ha, the highe:st
smoking rate, with neady two-thirds of men smoking. Today, apparently. one in eve,y three cigarettes
issmokedinOtina.
Globally, the consumpiion of manufactured cigarettes continues to rise steadily. G:insequently, the
death toll from the use of tobacco. currently esiimated to be 6 mUlion per year worldwide, is prro.icred
to rise to steeply (Figure 9.Z(b). This is inevitable because of the continuing popularity of cigarettes
and of the time lag between the taking up of smoking and the appearance of the sympiom.s of lung
cancer(Figure9.20a).
How has such a detrimental personal habit gained such a fOOl:hold worldwide? Why does it
continue to attract people? The answer must He panly in the impacr of the drug nkocine on the
central nervous system. Another is that people. particularly the young, have been (and are siill
being)persuadedthattheuseoftobaccoislinkedwithhighlydesirable,personalqualities,such
asadultsophistication,self-fulfilment,athleticprowessandsexualanracriveness
• Srudiesshowthatteenagersarescronglyinfluencedbytobaccoadvertising.
• Tcxlay, the multinational o:inglomerates that form the bulk of the tobaco:, industry have huge
budgets - in Asia, they are among the top ten advertisers.
In attemp1ing to deal with this health issue, the scale of industrial investment focused on this
product is a huge problem. lbere is a sound eo:inomic and moral case for continuous. powerful
•m•••L.
advenisingcampaignsthat moreeffectivelydiscouragetheuseofrobacco - despitethecostof
such initiatives. This is an ethical issue for all nations .
a) The 20-year time lag between smoking a nd the b) Prema ture dea th from tobacco use,
dia gnosis of lung cancer projectionsfor2000-2004and2025-2049
12 listtheeffectsof
tobacco smoke on the
cardiovascular system .
1250
.§200
i
I
Ia
150
100
so
I
• p,eventableifadu!tsquit(halvingglobal
c>garetteconsumptionby2020)
p,eventableif)<>tmgadultsdonotstart
(halvinggkibaluptakeolsmokingby2020)
othe,p,ematuredeatt.sfromtobacco-related
189
IJI Gaseousexchangeandsmoking
2 Theglobalcomumptionofmanufacturedcigarettes
continuestorisesteadilyandthehealthissuesgenerated
bycigarettesmokingareofepidemicproportions
Fig.1.1 worldwide
a) Suggestwhyyoungpeopletodaymaystillchooseto
a)OnacopyofFig.1 .1,drawlabellinesandusetheletters smoke cigarettes.
P, QandRtoindic.atethefollowingstructures b) Explainthechiefwaysthatcigarettesmokei1directly
P a blood vessel thatc.arriesdeoxygenatedblood harmful to the body.
Q astructurethatpreventsbackflowintoaventride c) ldentifywhycigarettesmoking,oncebegun, is addictive.
R abloodvesselthatc.arriesbloodathighpressure [3]
190
Examination style questions I
3 Scientists at the Tibet Institute of Medical Sciences in Lhasa 4 Fig 5.1 shows a drawing made from an electron micrograph
investigated differences between adult Tibetans who had of a cell from the ciliated epithelium of the bronchus
lived inlhasa{altitude3658m}alltheirlivesandildultHan
Chineseresidenuwhohadlivedthereforabout8years. The
Tibetam and the Han Chinese exercised at maximum effort
andvariousaspectsofthefrbreathingweremeasured
Someoftheresultsareshowninlable3.1 .
Tabl e 3.1
minutevclume/dm' min'
oxygenupt~ke/cm 3 kg' min'
• Minutevolume.Thisisthevolumeofairbreathedin
during one minute
• Oxygenuptake.Thisisthevolumeofoxygenabsorbed
intothebloodduringoneminute.ltisexpressedperkg
ofbodymass
The researchers observed that
• thegreaterminutevolumeofthenativeTibetansresulted
fromagreatertidalvolume
• the tidal volumes of the Tibetam !.hawed a positive
correlation with their vital capacity measurements
• theHanChinesehadlowervaluesforbothtidalvolume
and vital capacity.
a) StatewhatismeantbythetermtidaJvolume. [1)
b) Suggeo;twhytheresearchersalsomeasuredthevital Fig.5.1
capacityofthepeopleinthestudy. [2]
c) Explain how the minute volume at res t would be a) Copyandcompletethetablebelowby11,,-itingthe
determined [2) appropriateletterfromthediagramtoindicatethe
d) Suggeo;ttwodifferences inthestru cture ofthelungs structurethatcarriesouteachofthefunctionslisted
thatmayaccountforthegreateroxygenuptakebythe Thefirstonehasbeencompletedforyou (5)
TibetansshowninTable3.1 [2)
e ) When people who have lived all their lives at low altitude
facilitated diffusion of glucose
gotoaplac:eathighaltitude,suchaslhasa,theyare
oftenbreathless,lackenergyand~fferfromaltitude
sickness. However, with time, they often acclimalile to .,..,obKrespiratkm
the high altitude
In another study, researchers found that the red blood cell ~ ,iteoltfansoiption
count increasesin~chpeoplebyabout30%overseveral
-'"
Explainwhytheredbloodc:ellcountincreasessomuch
whenpeoplevisitplacesathighaltitude [2]
p;,c:kagespmteim into~mes
191
AS Level
1 Q Infectious disease
The infe ctious diseases studi edinthi s topicarecausedby sH ua l activity.Anund e rsta ndingofthe biology ofthe path ogen
pathog e ns that u e tra ns mitted from o ne hum a n host to a nothe r. a nd its mod e of tra nsmi ssion is essentia l if the disease is to be
So me, like Plasmo dium tha t causes ma la ri a, ue tra ns mitted by controlled a nd ultimat ely preve nted.
vectors;othe rs ue tra ns mittedthroughwater a ndfoodorduring
\/v'hilemany infectious By the end of this section you should be able to:
diseases have been
!.lKcessfullycontrolledin a) define the te rmd iseaseandexplainthedifferencebetweenaninlectiousdiseaseandanon·
50mepartsoftheworld, inlectiousdiseue (sickle ce llanae miaandlungcance r)
many people worldwide b ) state the name andtype ofcausativeorganism{pathogen)of e achofthefollowingd isea ses:
arestillatriskofthese cholera, ma laria, tuberculosis {TB), HIV/AIDS, smallpox and measles
diseases c) explain how chole ra, measles, malaria, TB and HIV/AIDS are transmitted
d) d iscussthebiological,socialandeconomiclactorsthatneedtobeconsideredintheprevention
andcontrolofchole ra,measles,mal ari a,TBandHIV/AIDS
e ) discussthe fa<lorsthatinflue nce the globalpatternsofdirtributionofmaluia, TBandHIV/AIOS
andusesstheimportanceofthesedi,easesworldwide
Disease: an abnormal
What are 'health ' and 'disease'?
condition affecting an By d isease we mean '3n unhealthy condition of the body'. lbere are many d ifferent diseases but
organism,whichreduces good health is more th3n the absence o f the h3rmful effects of disease-causing organisms. We have
the effectiveness of the alre3dyseentha! iJJne:ssmaybecausedbypoorlifescylechoicesorunfavou rableenvironmenml
functiomoftheorganism conditions,suchaslungcancerduerothesmokingofdgareaes(page184). Diseases of this
No n-infectious d iseas e type 3re called non-conununicable or n o n -infec tio u s dis eas es . They include conditions such as
adiseasewithacause chronic obstruaive pulmonary diseases (COPD) 3nd cardiova5Cular disease (CVD). discussed in
other than a pathogen, Topic 9. Non-infeaious disease m3y 3Jso be caused by malnutrition. In herited or genetic disorders
includinggeoeticdi1,0rders suchassicklecellanaemia3reincludedhere,too.
{suchassicklecellanaemia)
andlungcancer(linked
to smoking and other Infect ious disease
environmental factors} In this topic the focus is on dise:ises that can be 'cau ght" - sometimes described as conmgious
or communicable. lbese 3re infec tious diseases . They are caused by invading organisms living
191
10.llnfectiousdiseases l
bacterial disease
e.g.tube1Culosls
geoeticdiseas.es
e.g.haemopt,;lia.
sickle-cell anaemia
e.g.breastcaocer.lungcancer.
duetouncontrolleddMsionof
cells. causingtumou"
prions as disease agents
e.g.Creutzfeldt- Jacobdisease{C JO) allerg iesaOOauto-immunediseases - ove1Teaction
ofou,immur,esystemcancauseunpleasanlOf
dangerouse illr,e,ss.e .g.asthma
Figure10.1Typesofhumandisease
The pathogens that cause some infectious diseases exist permanently in a particular region or
Infecti o us di sease· population. Such a disease is said to be e nde m ic for it has a constant presence. Ma lar l.-. is an
adiseasecausedbya example - a serious threat to humans living in pans of Africa and other tropical and sub-tropical
pathogen that can be regionsintheworld.
transmitted from one host
On the other hand. e plde m lcs are diseases that appear suddenly and then spread rapidly in
organism to another.
a specific area or within a particular population group for a time. Cases of food poisoning are
Pathogen :abiological e xamples .
agent{suchasavirus, A p a nde m ic d L-.casc is an epidemic that spreads far more widely - typically throughout a
bacterium,fungusor whole country. a whole continent or throughout the whale world. It affects a huge number of
protoctist)thatcauses
people.lnthetwentiethceruuryseveralpandemicsofinfluenza('flu)occurred.Forexample,the
disease.A pathogen
'flu pandemic that occurred in 1918 was responsible for 10 million deaths worldwide
ca~nghumandiseases
In the preventio n o f infectio us dis e ase, knowledge of the biology of the pathogen is
willhave,aspartofits
structure,proteinsthat essential. So the fight against disease involves the work d microbiologists as well as the vigilance
aredifferentfromthoseof o fthehealthprofessionals. lnthistopic ,s ixspecilicinfectiousdiseasesofhumansareexamined,
thehumanhostandare most of which continue to present majco- threats to life. There are reasons why this is so in the case
therefore antigens of particular diseases but four general points are significant.
• We live in a 'global village' world with speedy, worldwide travel facilities available to some. A
person. infected on one COfltinent. may travel to 3nother within hours - before any symptoms
have appeared.
• The human population is in a phase in which it is increasing more and more rapidly
(Figure 10.2). In ever-increasing numbers. people congregate together - many in mega-cities.
Pathogens do not have far to travel!
• The work of health professionals is most effective where there is social stability. the absence of
war and an equitable division of wealth. Unfortunately, these conditions are absent from many
• As natural resistance to pathogens develops 3nc] drugs to overcome them are found. so
pathogensthemselvesevolvegreaterinfectivity thereisacontinuing'armsrace"between
pathogensand h oM s
193
ll:J lnfectiousdisease
Accidents
N«indudedinanydthe
typesofill-healthlistedin
Figure 10.1 is damage
duetoaccidents,suchas
broken bones. A bone
fracrureisnocadisease
butitiscenainlyan
unheahhyconditio nd 1000 yearsago -----* 265
___ I
(O¥er100000)'earsago)
Figure 10.2 The changing pattem of the estimated world human populat>on
194
10.llnfectiousdiseases l
__ .,.
(Figure 10.4).
--
\.1bliodloffae~
M>dreproduceln
,,..;;-~-;··.
....../ ~Na•ioo, Crlon;\.
---- t-• crm,
.............. ',~J
""' ./
\ fl,O
.
H,O i
~~ol)'teklls
Figun,10.4Theactionofthecholeratoxin
Treatment of cholera
About SO per cem of untre3ted rose,, of cholera 3re fatal but, when properly treated, generally
2 In cholera, toxic protein
less th,rn I per cent of people are likely to die from this very unpleasant d isease. A cholera
triggers the exit of
patient requires immediate oral (by mouth) 3dministration of 3 dilute solution of ions and glucose,
chloride ions from the
presentinarehydrationpack(Figure!0.5),inordertoreplacethefluidsandionslostfromthe
epithelium cells into the
lumenofthegut.How body. The presence of glucose aids uptake of ions and ccmpensates for energy lo.ss. This solution
does this movement prevents dehydration 3nd restores the osmotic balance of the blood and tissue fluid. This might
of ions differ from seem a simple process to people in the developed world but, in those pl3ces where cholera is
the movement of endemic, conditions may be very different. Many who contract the disease are weakened by
waterthatfollow.;ion shortage of food. Their drinking water may be of dubious purity 3nd have to be carried some
transport7 distance. The boiling of water makes it safe to use in the rehydration fluid but this requires fuel
that may be scarce.
Very severely dehydrated patients require the administration of intravenous fluids. An additional
treatment, the taking of the antibiotics streptomycin or tetracycline, wUJ assist in ridding the gut of
the cholera cells. However,thesedrugs3reoflinleuseunlessfluid,ions andglucosereplacement
has succeeded in restoring normal body fluid composition. An oral vaccine ag3inst cholera now
existsbutitisnoteffectiveag:,inst3llstrains.
195
ll:J lnfectiousdisease
Prevention of cholera
O>olern has been eradicated where there is effeaive sanitation. In the longer term, the disease can
bevinuallyeliminatedbysatisfacroryprocessingof....,ageandthepurificationofdrinkingwater,
including a chlorination phase. In endemic areas, the spreading of infection between households
may be prevented by the boiling of drinking water. lne faecal contamination of foods by flies must
also be prevented.
However. where populations have been displaced into inade<juate overcrowded refugee camps
or have migrated to crowded urban slums where dean water and sanitation are nOI: available,
cholera remains a threat to life (Figure 10.6). Malnourished children and those living with HIV are
at greatest risk.
Figure10.5Anoralnehyd,at;oo
packfortnl'treatmentofcholera
Figure10.6An,flJ91"camp-idealrnnditionsforchole,a
Table10.1Chole,a-tnl'profile
196
10.llnfectiousdiseases l
Malaria
Malarfa is the most important of all insect-borne disea..-s. posing a threat to 2400 million humans
in 90 countries - 40 per cent of the world"s population, in fact (Figure 10.7). About 80 per cent of
the world"s malaria cases are found in Africa south of the Sahara . Here, some 90 per cent of the
fatalitiesduetothediseaseoccur. ltisestimatedthataround400miJJionpeopleareinfected,of
which 1.5 million (mostly children under 5 years) die each year. Malaria kills more people than any
other o:inununicable disease, with the =ception of tuberculosis.
MalariaiscausedbyPlnsmo<li,im,aprotoctist.whichistransminedfromaninfecredpersonto
another by blcod-sucking mosquitoes of the genus AH<Jpbeles. lhe mosquito is descri.bo:l as the
vt.'Clor - anorganismthattransfersadiseaseorparasitefromonehostorganismtoanother.Ofthefour
species of Plasmo<lium that cause malaria in humans, only one (P.falciparum) causes severe illnes.s.
lhe others species trigger milder infections that are rarely fatal. Insecticides. particularly DDT, had been
effective againsi mosquitoes in the paSI but there has been a resurgence of malaria since the 1970..
;
Figure10.7Worlddi1tributionofmalaria
Until 100 years ago, malaria was endemic in Europe and the USA, too. In Britain it was called •fen
ague" because it was common in conununitie.'l living near marshes and because d the periodic
shivering fits patients experienced . Marsh and fen drainage for agriculture changed the environmeru
so that it was no longer favourable to mosquitoes and their transmission of the parasite. This is one
reasonf "ts d"sappe ranee ·n some areas.
197
ll:J lnfectiousdisease
Figure10.8Mosquitoleedingandtt>.,t,ansmi,.;ionolmalaria
Control measures
Control measures currently applied include the following.
• lnterrup1ionofthemosquit0Hfecyclebyattackingthelarvalstagewhichisanair-breathing
aquatic animal. Swamps are drained and open water sprayed with oil (which forms a surface
film,blockingthelarva"sairtube).andwithinsecticide.
• Use of insecticides to kiJJ aduh mosquitoes on and around the buildings that humans occupy.
• Protection from mosquito bites by sleeping under insecticide-treated mosquito nets and by the
burning of mosquito coils.
• Use of drugs to kill the stages of Plasmodi"m found in the blood and liver of infected people.
• Use of drugs to kill Plasmodium as soon as it is introduced into a healthy person's blood by an
infected female mosquito whilst feeding.
198
10.llnfectiousdiseases l
Figure10.9Thedistribuhooofsicklecellt,aitinAfric:aconl!'fSanadvant.lge
Adecermined,.,,ar.:hforavaccine.beguninthel900s,ledroun.suocessfultrials. Nowthesituationis
more hopeful. A huge injection d funds frcm the Gates Founcbtion into this project occurral in 1994
and recently more has been pledged. lbe subise<juent, energetically pursued se:,rch for a vaccine that
will confer lifetime acrive immunity for treated humans has yielded several promising 'candidates'. One
is undergoing advanced trials. lbe malaria r=r.:h community remains cautious; the comp!= P3rasitic
lifestyle d Plasmo.lium makes it a difficult to eradicate and there have been many earlier set-backs. ln the
longcrterm.controlofmo.squitoe-sbythecarefoluseofselectiveinsecticideswiUbeneceS&1ry. as,vell as
the development of powerful vaccines, before this very well-adapted malarial p:irasite is ove=me.
Table10.2Malafia-theprofile
Active immun it y
Pathoge n Pa thoge n taxonomy Pfasmodium. a prntoc:tist. with four species. of which P. fa/opdflim
immunity resulting from
isthe m05ldangerous
expornretoanantigen
During the subsequent Via the blood meal, of the fem ale Anophele, mosquito
1mmunere1ponse, Diseaoe Incubation and venue Pfasmodium enters liver cell,. reproduces asexually. then parasites
antibodies are produced pararneten are released into the patient~ bloo d stream and <'f11er red blood
byplasmacells andthe cellsimmedia tely.Acycle of ,edbloodcell iovasion. ,eproduction
body makes memory cel ls arid ,elease(withtoxiros)isthen,epeated
th atprovideongoinglong- Signs.symptoms and A cydeol ve,yhightemperatu, es,headache,nausea ariden!a,ged
term immunity. Thereisa di ag nosis tende,spleeneve,y48-72hours. followedby lily
delaybeforethe immune
responsei:scomplete,so A comb;nation of anti-malarial drugs
immunity ta kes someday, Prognosis Most patients imprnvewithin4 8hoursa fte,initiatioooftreatmenl
to build up andbecomefever-freea fte,96hours
lnfectionbyPfa/dparomcaus,,saf<lfmofmalaliawithahigh
mortality,a te ifuntreated
199
ll:J lnfectiousdisease
Tuberculosis (TB)
Tu Derculo,;is (TB), a major. worldwide public health problem of long standing, is caused by a
b acillus (rod-shaped bacterium) called Mycobacterium mberculosis (Figure 10.10). Viewed under
the microscope after staining by the Ziehl- Nielsen technique. these =lls appear bright red. This red
colouration is unique to bacreria of the genus Mycobacterium and is due to wax and Olher lipids in
their walls. This is how the pathogen is identified in infected patiems
Figure10.10Hectro11m,aograph
olacolonyol/v!rrobaaerium How tuberculosis spreads
ruberculosi.-{x25000)
People with pulmonary TB have a persistent cough. The drople1s they inevitably spread in
the air are infected with Jive Mycobacteri11 m (Figure 10.11). Tuberculosis is chiefly spread by
droplet infection in this way. Because of their lipid-rich cell walls, the bacilli are protected
from drying out so the pathogen may survive for many months in the air and the dust of
homes . This is anO!her source of infection. Overcrowded and iJJ -ventilated living conditions
are especially favourable for the transmission of the infecrion. However, it still requires quite
prolonged contacr with a viable source before people will succumb, because the bacterium is
notstronglyinfecrious .
A bovine form of TB occurs in cattle and the bacillus can enter the milk . Unpasteurised milk
from infected cows is anorher potential source of infection. Jn some rural communities especially,
this type of milk may be consumed by both adults and children. Today. in some countries. milk
is supplied from 'tuberculin-tested ' cows that are certified free of Mycobacteri,m,. However, TB
may stiJJ be found in dairy herds. This is because of a 'reservo ir' of the bacterium that exists in
w ild animals that have chan= contacrs with the herds. Treatment of milk by pasteurisation is a
matCJr control mechanism as this hem treatment kills the bacteria without affecting the quality
o f the milk .
Figure10.110rnpletinfectioo
How TB develops
Onceinsidethelungs,thebacteriaareengulfedbymacrophagesinthealveoliandbronchioles.
If the person is in good health these white blood =lls (with the help d the T-lymphocytes which
migrate in from lymph nodes - page 159) kilJ the pathogen. Alternatively, and panicularly with
strains of Mycobacterium that are more infecrious, the pathogens may remain alive within the
macrophage, although localised and effecrively controlled by the immune system. This =plains
w hy it is that not all infecrions with M. tuberculosis result in TB - many people show no sympioms.
Jn facr. about one-third of the world"s population is infecred with
M. l!<bercu/osis, but only S--10 per =m becomes sick or infectious
at somestageduringtheirlife.
However, if the patient is malnourished or is in inferior health with
aweakenedimmunesystem,achronicinfectionmaydevelop.typirnlly
w ithinthelw,gs. Cavitiesappearasbacreriade:stroythelungtissues
Blood vessels are broken down and fluid o:illects. The strucrural
damage to the lungs can be seen X-ray examination (Figure 10.12).
If the condition is not treated prompily. the pathogen may be
carried in the bloodstream and lead to TB in almost any pan of
the body. For example , the meninges of the brain , bone tissue, the
lymph glands, the liver. the =mral nervous system, the kidneys
or the geniml organs can all be atmcked. Generally, patients show
loss of appetite, loss d weight, excessive sweating and a decline in
physical activity.
200
10.llnfectiousdiseases l
Treatment of TB
Today, TB is treamble but it is still a 'killer' disease if not diagnosed early on in the infection
Amongst those with the disease of HIV/AIDS , TB is the leading cause of death .
Since TB is infectious, on confirmation of the diagnosis of a case of aaive tuberculosis, all
comacrs of the patient n=d to be traced and screened by a community public health team.
Infectious patients require isolation and treatment with specific antibiO{ics until they cease to cough
up viable bacilli. Then, antibiOl:ics continue to be administered to the patient, now back in the
community, until there is evidence the infection has been eradicated from the body
Unfortunately, microorganism.,developresisrnncetoantibiocicsanda1herdrugsusedagains1
them, with time. In the case of TB, this has already happened. Patiems are now treated with several
antibi(){ics simultaneously because of the emergence d multidrug-resistant TB (MDR-TB). This is
particularly in the case of A!Ds patients . MOR-TB is especially common in the USA and South-East
Asia (seeTablelO.J).
Prevention of TB
A vaccine against tuberculosis. the BacUJus Calmene---Gu&in (BCG) vaccine, is prepared from a
strain of anenuated (weakened) live bovine tuberculosis bacillus, Mycobacte,1um bods. It is most
often used to prevent the spread d TB among children.
A tiny quamity of inoculum is injected under the skin of the upper arm . Before vaccination
takes place. people are tested for existing immunity. since these people will re:icr unfavourably
to the vaccine (the Haef test, Figure 10. lJ). Only those with no re:icrion may receive the vaccine.
Immunity typically lasts for at least 15 years - longer if the individual is re--=posed to TB bacilli ,
for example, by accidental encounter with it . At best, the BCG vaccine is effective in preventing
tuberculosis in a majco-ity of people for a period of 15 years. However, immunity wanes with time
anddisappearsentirelyfromtheelderly.
Figure10.13HaeltestandaTBvaccination
201
ll:J lnfectiousdisease
How effective the vaccine- is varies according to geography. Trials of ECG in two Slates in the USA
were Oflly 14 per cem effective. Another trial showed no protective effect. In the UK the vaccine
is 80 per cem effective. How BCG vaccination is applied in different countries is reviewed in
T3ble10.J
Colffltry Vaccinationpolicy/re«>mmendedfor
Unite d Kingdom Babies and child,en unde, 16 yem with pa,ents o, g,an~a,ents born in counhes with
high,etesofTll.or"tierebom inonethemSl'M.'S
Thosewhohavebeenindosecontac:twithaTBpatient
Thoseundettheageof35whoseor:rnpatkmputsthematriskofexposuretoTll
lntmdtKedin1948 - thefirstcountrie'.loutsideEuropetodoso
Death s fromTBinEngla nd a ndWa les, 1900-2000 The changing pattern of the disease
uptumpo,,;iilyattriiutable
decline attributable to
to globalisation of There is evidence that m was present in some of the
impH1Yementsin e,irliest human communities and it h3S persisted 3S a
housingprnvisioo populatkmma,ements.
and appearance of m3jor threat ro he:ihh where vulnerable people lived
anddtflforthepoo,est
people in
bacill i resistenttoavailable in crowded conditiOfls. In the ninet=nth century.
society drn<o>
when !he TB bacillus W3S first identified, Ofle in seven
of all de3ths among Europeans was due to TB.
Tochy.inmostdevelopcdcountries!hisdiseaseis
relativelyrare - intheUKforexample. in 2007the
national average was about 15 cases per 100 OOO of
first anti-TB drug the population. In these countries, the reduaion in
treatmentsdisc...,,.,.,.J
TB!hroughoutmostofthepastcenru,yrulsbecndue
BCGimmunisatkm tosteadUyimprovinglivingcOflditioru.particul3rly
•i 30 prog,ammeswklely
adopted
inhousinganddiet(Figure 10.14). However,today
.,
1 20
thereisevidenceofaresurgenceof!hedise:isedueto
inward migration of new dti7.ens, to the globalisatiOfl
of travel and to !he emergence of multidrug-resismru
TB(MDR-IB).
Globally, almost 14 milHon people have active
TB, with approaching 2 million de:iths due to the
bacillus annually. lndevelopingcountries withahigh
incidence of this dise:ise it primarily affects young
Figure 10.14 How ,;or:iaj, economic ar>d medical factors~. influenced the oa:urrmce adults. In developed countries where TB had ceased
ofTilindev,,lopedcoootries tobeamajorhe:ihhthreat,!herisinginddence
ofthediseaseislargelyamongtho.sedescribedas
immunocomprcmised (page 205). particul3rly those
with HIV/ AIDS, 3nd among new migrants from
devdopingcountries - spcciallywhere!heyarenow
livinginpoorerhousinginlargedties. such3SNew
202
10.llnfectiousdiseases l
Table10.STubercuk>sis theprofile
203
ll:J lnfectiousdisease
HIV/AIDS
Amo.Immune deficiency syndro m e (AIDS) is a disease caused by the human immunodeficiency
virus(III\', Figurel0.15). Vlrusc~aredisease--causingagents,rathcrthan 'organisms'.
lnkc tion wilh IIIV is only possible through
contaawit:hblocxlorbo.Jyfluidsofinfectedpeople,
suchasmayoccurduringsexualintercourse.
Antib Kltbarenoteffective
§:~~
tr;om:rlpl.lse
sharingofhypoclennicneedlesbyintravenousdrug
users,andduringpregnancy,labour.delh=yand
breastfeedingofanewbombaby.Also,blocxl
trarufusionsandorgantransplantswilltransmitHN
against viruses.Wha t antibiotics
.....
(eozym<")
.,_,
butdonorsarenowscreenedforHNinfea:ionin
-
areand"flythisissois most muntries. HN is not transferred by cont:1<1 with
exp!ainedonpages210--16 salivaonadrinkingglass,norbysharingatowel,for
example. Nor does the female mosquito trarumi! HN
(pro\'-"n) whenfeedingonhumanblocxl.
Antigen:aprotein(normi!lly The spread of HN and the eventual onset of
Figure10.15Thestructureofthehuman
~some rarix>hydrate'i and immunodeficiencyvirus(HN) AIDSinpatientsareoutpacingthecurrentefforts
"""'""""""'"'"'
canactasantigens)that
isrecognisedbythebody
of scientists and doctors to prevent them. 1be
World Health Organisation WHO records the current Slate of this pandemic (Figure 10.16).
204
10.llnfectiousdiseases l
figure10.17HWinfechooola"tiitebloodcell
HN is a virus belonging to a group known as retroviruses. Rc lrovlruscs are RNA viruses with a
unique "infection process'. On entering the cytoplasm of a host cell the virus RNA is translated into
DNA which then attaches to that of a chromosome in the host's nucleus (Figure 10.18). The steps ro
this are as follows .
Inside the lymphocyte. the RNA strands and an enzyme rnlled reverse transcripiase are released
from the core of the virus. Then, using the viral RNA as the templace, a DNA copy is formed by
the action of the reverse transcripw.se. (Remember, RNA is single stranded but DNA is a double-
stranded molecule, page 112).
This DNA enters the nucleus of the lymphocyte and anaches itself to a chromosome. It becomes
a permanent pan of the host cell's geJX>flle. It is known as a provirus. After an initial , mild form of
AIDS (a brief 'flu -like illness within a few weeks, or possibly the developmeru of a rash or swoll=
glands) the provirus remains dormant (called the latef>C)· period).
~--
2DNAcopylomled
fl'YefSetramcrlptase
=~
3 RNA<traffin>ml'l'l.'d
,_..,.,
,uamcrl!)llonol
OOAstrandbyaalol!
olONApol)'mera<e
50NA-<traffi
~7~~~
_,,_
ilnodells<Ahosl
("'tem1'1rus)
Figure10.18Hw,H1Vbecomespartolthewhitebkxxlceltsgenome
However, at some later time the viral DNA replicates, leading to the produaiOfl of new viruses
which then invade and kill other lymph0<.)"te5 (Figure 10.19). It is not dear whether infected
T-lymphoq"tes are killed by the virus they harbour or by the aaions of the patient"s immune
system against these infected cells. However, AIDS-related symptoms follow, mused by pathogens
tha! are around us all the time. These are normally resisted by a healthy immune S)·stem but,
without it. the lxxly cannot effectively resist (Figure 10.20). We say the patient has now become
immunocompron ·scd .
205
ll:J lnfectiousdisease
~
,.-n
. ,_
i'~•<om
,
oppo rtunisti c infecti on
{e.9.P™'oo,or,ia)
takesmer - pat>efn~
i=une,ys1emnow
·~r·
,oogefor
- ~IV=copies"°', mm'
messer,ger
RNA copy
to,;bosomes
-"'
HNonhost
-· l
sigr,fc.aMly
T-l;rnphocytes
l -
seVl'<l'T-l;rnphocyte
deple!ioo
i
,h-=•
~'"
Figure10.19ActiVilhoooltheHIVgenomeandtheproductionofnewHN Figure 10.20TheprofileofanAIDSinfection
206
10.llnfectiousdiseases l
Table10.6ThepreventionandcontrolofHW/AID5
Effectiveeducabonprogrammessothatthe Contacttfacing.whe,eaperwndiagnosed a,
vulr.eraMeunderstand thecauseandeffects HIV-posrtiveiswi lli ngandabletoidentifythose
ofHWinlection.andthebeststepsto,ema in "flohavebeenpotatriskbyrnntact
heak:hy.whether0<oottheyarelrterate
Encouragingla ithful1>esstoonepartner
Condomsbeingmadeffeelyavailabletothe fundingof testingofthesexually-active
sexually active population populationtoidentifyHIV-posrtivepeople(if
5 Suggest what features SterileneedlesbeingmadeTI"eelyavailabH'to r,eoplewill acceptthisirwasionofthei,pfivacy)
oftheHIVvirusandthe intravenoosdrugusers fundingandsupplyofthedrugsthatprevent
disease it causes create thereplKationof thevirus insidehmtcelis
particular problems Biolog ica l Thepractic:eof"safesex"bytheuseof Drug therapy
forthepeopleandthe coodomstopreventthetfar,smissioothmugh Resea,chonthedevelopmentofavaccine
economiesofless-
infectedblood0<semen Replacementofbreastfeedingbybott~
developedcountries Sc,eeningofblooddonorsfo,HIV.,othat feedingofinfa nt,ofHIV·positivemothers.
!.UChasZimbabweor infectedbloodisnottakenaOOused andothe,WHOstfategiesl0<helpingHW·
Zambia? infected mothers
Table10.7H!V/AID5-theproMe
207
ll:J lnfectiousdisease
Measles
Measles is a highly infectious disease that people of any age can get. It is most commonly
contracted by children aged between one and four years. Unlikeotherchildlxxxldisease-s. such as
mumps and rubella. it can generate severe complications. Measles is a virus disease entirely specific
to humans and with a long history. For example, there is a temh--century record of it being 'more
dreaded tba>1small/)OX. In fact , in the past most people contracted measles. The incidence of this
infectionandthedangersitposedwereabruptlydiminishedonlyasrccentlyasthe 196().s.Acheap
and reliable vaccine became available then but it was effective only when taken up by 95 per cent
of the population. This is often the case in the developed world (but nOI: always - see below)
Today,over95percemoffatalitiesduetorneas.lesoccurinlcss-developedcountrieswherehealth
infrastructures are poor. Measles is also a major killer of those in an advanced stage of HN/ AIDS .
Pa ssive immunity
immunity involving the
transfer of antibodies
{alreadymadeinthebody
of another organism or
invitro}intothebody
Figure 10.21 Number of measles cases in the USA. 1944-2007
wherethey'Aillbindto
theirspecificantigenifitis
A combined vaccine against three common childhood illness, referred to as the MMR vaccine.
present.Thisgivesimtant
became available in 1971. The MMR vaccine is made by using live, attenuated (weakened) strains
immunity but does not
lead to the development of the measles, mumps and rubella viruses . Infants in the UK, for example. are now offered a first
ofmemorycells,'iOlhe MMR injection aged lS--24 months and then receive a booster injection between 3 and S years' of
immunityonlylast1fora age. (A baby under 6 momh.s is protected by antibodies received from the mother. via the placenta,
few weeks whilst ;,, ,<tero; conferring what is known as ' p assh'C immunily'.) The success of vaccines has
made measles a rare disease, at leasi where the vaccination regime has been adhered to.
208
10.llnfectiousdiseases l
Table10.8Measles-theprofile
fstimated 900 OOO death, from measles {,008). in co,mtdes with low
a nd mortality incomesandlimrtedhealthinlra,t,uctu,es
Why did some parents amfidently blame !!U!Rfor their child's co'1dition?
Autism, when it arises, does so at about 18 months. %en this 0ct.1.1rs soon after the MMR
inoculation. it is inevitable that there mar "ppear ro be a connection. In fact. no stud)' has
produced evidence that MMR causes autism . Major investigations and follow-up studies have
beenundertakenwithlargesamplesofchildreninAustralia,inFinland,andinJapan. In
communities where autism appears to be on the increase, the increase began before MMR was
introduced,andtheineidenceofautismcontinuedtoriseevenwhenthenumbersofchildren
7 The coincidence of receiving MMR reached a plateau. There is no doul:x in the minds of health authorities that
autism onset and vacdnationisthesafestopiionforchildrenandtheirparents.
MMRadministratioo The measles vaccine is nOI universallr available-, there are several less developed countries
maybe described as where manr people are nOI vaccinated. This is despite the fact the disease is targeted by the
circumstantial evidence WHO in an expanded programme of immunisation. It is estimated there mar be up to 900 OOO
1/v'hatpart,ifany,can
measles-related deaths annuallr - man)' HIV/ A!Ds-rdated. Unfortunatel)', some children who
itplayin1cientific
recovcrareleftwithserious,permanentcomplicatioru
discovery?
209
ll:J lnfectiousdisease
10.2 Antibiotics
Introducing antibiotics
Antibiotics were discovered relativdy recently - with a chance observation in 1928. By the 1950s,
it became possible to treat patients with courses of antibiotics that cured many bacterial infectiot1S.
More recently. the excessive use of antibiotics has generated problems, a., we shall see later.
AntibioticsarenaturaUyocrurringchemicalsubstancesobtainedmainlyfromcertainfungiand
bacteria commonly found in the soil. When antibiotics are present in low concentrations they inhibit
the growth d other microorganisms or cause their outright death, a., demonstrated in Figure 10.23.
lbediscovery. isolation and development dthe firstantibictic, penicillin, was not an easy task. Ot rook
from1929untill944.) Sincethen.overfourhundreddifferentantibioticshavebeenisolatedandtested.Of
these,onlyabout50haveprovedtobenon-toxictopatients. lheseant1bicticshaveachievedwideusage
lhoseeffectiveagainst a wide ranged pathogenic bacteria are caUed broad-spc.-ctrum antibiotb. lbese
include chlornmphenicol . Others, including .streptomycin. are effective agains{ a limited range d bacteria .
210
10.lAntibiotics .
Toak"""'ns.pec,e<;oibacteria
gro,,ingoonu!Jieotaga,wa,
adledamasrrnQwi!heach'arn,'
imp;egnatedwi!had:ffe,..,,t
antibiotic(coloo,coded).Ttlen
~:!i~::~~t
(CJWOl,ite)th,e,ei:,.,.,,dence
t~s,~~h~~=~001
antibiotics(e.g. CM,A)
thaoroo!hers(e.g.S,I)
kOO'Mlsp<'Ol"ol
:::~~
medium
:'nawhe"'
haYebem!uled
Oiffefenlaotibiotics,..,.comair,edin!heannsolthe
masrriog,sothat"'m~Mty to manyaoti biotiamaybe
lested5'll'Uta<le00Sly.
Figure10.231nvestigatir,gsensitivitytoantibiotics
211
ll:J lnfectiousdisease
Today, the issue of bacterial resistance to amibiOlics in healthcare has become a major problem.
Aspathogenicbacteriadevelopedgenesforre.sislancetoantibioticaction.s.differentantibiotics
were used in response. The inevitable outcome of this is that pathogenic bacteria slowly acquire
mult ip le resista n ce (Figure 10.25). For =ample, a strain of StapbJofococcus a"reus has acquired
resistance to a range of amibiOlics including methicillin. Now. so-called methicillin-resistant
S1apby/,x;or;cus ao,rew (MRSA) is referred to as a "hospital superbug' because of the ham,
itspresencehasinflictedintheseplaces.(Actually,"superbugs"arefoundeverywhereinthe
community, not just in hospitals.) MRSA poses the greatest threat to patients who have undergone
surgery. With cases of MRSA to treat. the intravenous antibiotic called vanomycin is prescribed. but
recently there have been cases of panial resistance to this drug, too (Figure 10.26).
Similarly, a strain of the bacterium Oosrridium difficile is now resistant to all buc two antibiotics.
This bacterium is a natural componem of our gut 'microflora·. It is only when C. difficile"s activities
are no longer suppressed by the surrounding, huge, beneficial (i.e. 'friendly') gut flora that it may
multiply to life-threatening numbers and then may trigger toxic damage to the colon.
212
10.2Antibiotics .
~thogef,inv~..ld trigrJN,~
lnoomaon(orin•petor farm..-.imak)
9 Explain·
a whydoctOfSasi::
patientstoensure
that they complete
t~rcourseof
antibioti cs fully
::.::r:lt!•noll>H
=~~f~NMI
antibio!O::e.g. meth icilin)
=i'::::::::.·
owruwandfmC/WolammioticsaNtelhe
pefeC1 bf-iinggrO<rd for resi<lant ~mri.o
llllhelongterm.lhPdrug,indusuyfau-s
:~::,:t!'oc"'!"'~7':!:::..
!Uf
~lopmentof muhlple-r.sistanl
sraphylrxomiJaur•us. 1992-200 1
('11,ollnfectedsarnplesreceivedlnJ)itl"dogylabs)
Fl,g...-•10.26The1nUNS1ngin::iden::eofMRSA
113
ll:J lnfectiousdisease
Figure10.27Fake-coloo,SEMsofSrilpliylococcus.wrros(xOOO)andC/osrridiumdiff,ci/e(x10000)
THETIMES1ThuMayMay 12014
year interval,
Figure10.28Thenumbe,ofnewantibioti<sdevelopedand
approvedinthepastthreedecades
214
10.lAntibiotics .
Inthissiruationthestrategytocombattheproblemincludes:
Geo,gegets
, _ _ ant,boobcsand
devek,psres,stant
bacteria in his gut
~::rn ~ ....____
gene,alcommunity.
:::resistant ""
I r--- ::~:f
Fertilise,o,wate, Resistant g e r m s s p r e a d &
containing a nimal faeces difectlytootherpa1,er,1,or
anddrug·resistantbac:teria indirectly on unclean hands
isu!l'donlooduops olheatthcareproviders ·
Figure10.29Howant1bioticresistar,cesp!l'ads
215
ll:J lnfectiousdisease
116
Examination style questions I
Examination style questions
1 a) i) Nametheorganismthatcausestuberculosi1{TB}. [1] 2 a) Movestopreventthespreadofinfectiou1disease11,ithina
ii) Explain how TB is transmitted from an infected penon local community have included
to an uninfected person [2] A an improved supply of wa ter, including sewage
The World Health Organisation (WHO) collects data on TB treatment
from its six different regiom as shown in Table 1.1. B insectcontrol
ln2003,itusedthesefigurestoestimate
C milkpasteurisation
• thetotalnumberofpeoplewiththedisease ineach
region Foreachofthesepreventativemeasures,describea
• thenumberofdeathsfmmTB. diseasetheyareeffectiveagainstandexplain1\tiy. (12]
Many of those who died from TB were also infected b) Whatfeaturesofthelifecylceofthepathogenthatcauses
with HI\/. malaria make the development of an effective vaccine
Tab le 1.1
especially difficult? [4]
c) Explain why malaria is endemic in tropical and 5Ub-trcpic.al
Number of deaths from regionsbutabsentorrareelsewhereintheworld [2]
Number of cases TB( includingTBdeaths in d) Whatisthesignificantdifferenc:ebetweenadisease
per100000 peopleinfectedwithHIV)
outbreakwhic:hisdassifiedasanepidemicandasa
WHO region population per100000population
pandem ic? [2]
[Total: 20)
217
AS Level
11 Immunity
An unde rsta nding o f th e immun e system shows how ce ll s a nd th e same path oge n. Phagocytosis is a mo re imm ediate non-spec ific
mol ec ul es functio n togethe r to protect th e body ag a inst in fectio us pa rt o f t he immun e system, w hil e th e action s o f lymphocytes
di seasesandh ow th e body is protectedfromfurth er infec tion by p rovideeffective de fenceaga inst specific pathoge ns.
The immune system has By the end of this section you should be able to:
non-specificand~ific
responses to pathogens a) state that phag ocytes (macrophages and n= trophil,) have the ir origin in bone marrow and
Auto-immunedi1ease1 descr ib e the irmode ofaction
aretherernltoffailures in b) descr ib e the modi!'< of action of 8-lymphocyt"' and T-lymphocyte<
the1ystemtodi1tinguish c) decr ib e and6plainthe significance ofthe incre...e inwhite bloodce llcountinhumanswit h
between self and non-self
d) explain th e meaning of the te rm immune response, making re fe re nce to the terms antige n, self
Responses to infecti on
Pathogens do not gain easyenayto thelxxly because:
• =remally, the keratinised protein o f the dead cells of the epidennis are tough and impervious
unlessbroken,cutordeeplyscratched
• internal s,ufaces, inpanicularthetrachea, bronchiandthe bronchio lesofthebreathingapparatus,
and the gut are all lined by m o ist epithelial tissue. These internal OOniers are protected by mucus,
by the aaiOfls d cilia removing the mucus, and some by digestive enzymes or strong acid (as in
the stomach).
However, these barriers are sometimes crossed by pathogens. Jn response. there are internal 'lines
o f defence'. Firstly. the body responds to localised damage (cuts and abrasions, for example) by
infbmmatlo n . If a blood vessel is ruptured then the blood clo ning mechanism is aaivated. Jn the
blood and tis.sue fluid, the immune system is triggered, which is covered in more detail shonly.
Inflammation
Inflammation is the initial, rapid, localised response the tissues make to damage, whether due to
a cut, scratch, bruising, or a deep wo und . We are quickly aware that the site d a cut or knock
(contusion) has become swollen, warm and painful. Inflammatio n is triggered by the damaged
cells themselves, which release 'alarm' c hemicals, including his ta mine and prosraglandins.
218
11.1 lhe immunesystem I
The initial outo:ime is that the volume of blood in the damaged area is increased, and
stemcellsinthebooemarmw
white blood cells and plasma accumulate outside the enbrged rnpillaries. Ultimately,
~
cells mig,atetothe!ymphnodes
tissuerepairisinitiated,also.
The blood circulation pbys a o:implex pan in the resistance to infection that will
result if microorganisms enter the body. The Increased blood flow retn0,..,s Loxlc
ph agocyticce lls '(I) produ cts released by invading microorganisms and by damaged cdls (Figure 11.J)
The white blood cells that accumubte at the site of tissue damage or invasion of
~::~t~:,~i
pathogensfallimotwofuncrionalgroupingsdependingontheirrolesinthedefence
against disease.
General phagocytk white cells engulf 'foreign" material. These are the
white cells nemro phils and the m acrophages . Neutrophils make up 6o per cent of all white
• high!yeffidentatengu\fing blood cells in the blood, but they are short-lived. Macrophages are the principal
bacte<iaeithe, inbloodor
'rubbish--collecring cells" found throughout the body tissues (Figure 11.1). lbese
intissuefluidsoutsidethe
cap illaries long -lived responses toinfecriondestroyany invading pathogen.
~:::hat ~
• surviYeoo!yafewdays ph agocyticce lls Other white blood cells called lymphocyu,s, produce the antibody reaction to
infecrion or invasion of foreign maner. This, the immune response, is triggered by and
directedtowardsspocificpathogens .
You can see some phagocytic white cells and a lyrnphocrce in the photomicrograph
of a blood smear, in Figure 8.2. You will probably have already seen them in a
~~:~~~~bbish~ole<ti~\@ prepared microscope slide of mammalian blood.
• leavethebloodcirrnlationandlte., , All these white blood cells originate by division d stem cells in the bone marrow
;nwaitintheliver,kdneys,spleen :• and in cenain cases they mature in the lymph nodes. On average, they spend only
and lungs · • about 10 per cent of their short lives in the blood, but all are o:iocemed with the body's
• always ready to respond to debris ~
proiection against infection
and invading cell,
The blood also delivers special proteins (complement proteins), which are
Figure11.1Thephagocyticcel!solthe acrivated by the presence of infection. Complement proteins enhance the work of white
body'sdefencemecl1anism blood cells in overcoming infections. &me trigger the lpis of invading microorganisms.
and others, the opsonins, bind to pathogenic bacteria and so increase phagoc)'{Osis.
Cells infecred with ,iruses produce proteins rnlled imcrfrrons . These bind to
neighbouring, healthy cells a nd trigger synthesis d antiviral proteins . Viral replication is
halted.
219
m 1mmunity
interferon
~:!.'%~
neutrophils eng,u~edbacteriumin'food
~;-!":n:7':
l>yopsonin:s
vac:oole' - !ysoso,neswilhhydrnlytic
enzyme,;fuse,c.>Usingdestroctioo
~ of bacterium
Figure 11.3 lnftammahoo at a damage site, the processes
220
11.1 lhe immunesystem I
has distinaive MHC antigens present on the cell surface membrane of mosi dour body cells.
Unless you have an identical twin. your MHC antigens are unique
Lymphocytes of our immune system have amigcn n,cq>lors thm recognise o ur own MHC
antigens and can tell them apart frcm any •foreign' antigens decected in the body. After all , it is
critically important that our own cells are nOI anacked by our immune system.
Se lf : the products of the body's r:mn genotype, which contain proteins {normally ~ §Orne
carbohydratesandothermarnimolerulescanactasantigens)thatdonottriggeranimmune
response in the body's own immune system. Inside the body that produced them, self proteins do
not act as antigen1 {and 10 do not 1timulate an immune respome} but, if introduced into another
body,theybecomenon-1;elf.
Figure11.4T·andll-lympho<.ytes
221
m 1mmunity
Figure 11 .!.Thestructureofan
antibody
2 On binding to the B-lymphoc:yte. the antigen is taken into the cytoplasm by endocyrosis, before
being =pressed on the cell surface membrane of the B-lymphoc:yte.
3 Meanwhile. T.ly mphoq·tes can only respond to antigens when presented on the surface
o f other cells. Phagoc:ytic cells of the body, induding macro phages, engulf antigens they
encounter. This may occur in the plasma, lymph or tismie Ouid. Once these antigens are taken up,
the macrophage presents them externally by attaching the antigen to their surface membrane
pr<>{eins. MHC antigens. This is called amlgc n p n,scnmtlo n by a macrophage.
4 T--cells come in contact with 1hese macrophages and briefly bind to them. The T-lymphocyte is
immediately activated . They become actfr:u cd helper T-ly mphocytes.
5 Activated helper T-lymphocytes now bind to B-lymphocyres with the same antigen expressed on
their cell surface membrane (step 2 above). The B-lymphocyte is activated by the binding of this
chemical. It is now an act h -a ted II-ly mphocyte.
6 Activaced B-lymphocytes immediately d ivide very rapidly by mitosis fanning a done d cells called
plas m a c ell... Plasma cells are packed with rough endoplasmic reticulum (RER). It is in these
organelles that the antibody is mass produced. The amilxxly is then exp:,ned from the plasma
cellbyexocytosis. lbeantibodiesarenormallyproducedinsuchnumbersthattheantigenis
over=me. lbe production d an activated B-lymphocyte, its rapid cell division to produce a done
of plasma ceUs andtheresultingprcductionofantilxxliesthatreacrwiththeantigen is called
clon a l selection . Sometimes several different antilxxlies reacr with the one antigen - this is
poly d o n a l selec tlo n .
222
11 .llheimmunesystem I
FiguN11.G Thesuge,s111¥11bodyprodu::tnl
113
m 1mmunity
7 After these antibodies have tackled the foreign matter and the disease threat is overcome,
Na tur a l immuni ty · theamibodicsdisappearfromthebloodandtissuefluid. Soroodothebulkofthespecific
immunity t hat is acquired B-lymphocytes and T-lymphocytes responsible for their formation.
bytheindividualasa However, rerrnin of these specifically activated B- and T-lymphocytes are retained in the body as
naturalpartoftheirlife m e mory cells. lbese are long-lived cells, in contrast to plasma cells and activated B-lymphocytes.
Thisindudesnatural
Memory cells make possible an early and effective response in the event of a re-infection of the
passive immunity following
body by the same antigen (Figure 11.7). This is the basis of natural immunity.
transfer of maternal
antibodies intoafetus
throughthepli!Centaand Memo,ycellsa,e
into anewbominfant in ,etainedinf;mphn.ode<
thefirstmilk(colO!trum} TheyaHowaqukl<and
ltal'iDindudesthenatural spec;fkres.pooseifthe
sameaotigen,eappears
active immunity that
follows natural infection
by a pathogen involving
the production of memory
cells{forexample,natural
infection1\ithchid:enpox,
giving long-term protection
against this virus)
Figure 11.7Theprofi\eofantibodyprodtKtiononinfectionand,e-inferuon
2 ldentifywhereantigens It is now helpful to summarise the comp]= roles of B- and T-lymphocytes in the immune system
and antibodies maybe (Figure\1.8)
found in the body.
3 Remindyour5elfofthe
appearance of the types
of white blood cells
The significance of white blood cell counts
observed in a blood A normal white blood cell couru is between 4500 to 10000 cells per pl, but during active infections
smear preparation in the body the number rises and may be considerably higher. Given the roles of white blood cells
Seeforexample,the when in the presence of pathogens within the body, this relationship is of no surprise. Another
photomicrograph in cause of raised white blood cell count is leukaemia - a cancer of the bone ma!TOW cells. The seem
Figure8.2{page1S3}. cells from which white blood cells are normally formed proliferate but fail to differentiate. Here the
Compare thi1 image numbers can be very high indeed. but with a variety of abnormal cells present.
withaprepared!lideof
a human blood smear
sothatyouarefamiliill"
with the appearance of
Myasthenia gravis - a failure to distinguish
phagocyticwhiteblood
cells and lymphocytes 'self' from 'non-self'
Make a fully Myastheniagravisisanusualauto-immunedisorderinwhichspecificamibodiesanackanddestroy
annotated drawing of the acetykholine receptors at the post-synaptic junctions, thereby inhibiting the effect of the
representative white neurotransmitter acetykholine ( page 319). This disease is one of the Jess common auto-immune
bloodcell1thatmake1
disorders. affecting about two people in every 10000 of the population. It causes muscle weakness.
clear the differences
The effects of the disease may be localised in the body, involving just the eye muscles for example.
between them.
Alternatively, many muscle systems may be affected, leaving the sufferer susceptible to general
fatigue. In the presence o f this condition, muscles weaken during periods of activity but improve
with resting. Some patierus with myaschenia gravis are found to have a tumour of the thymus gland.
These are often benign, but the gland is then removed as a precaution, and typically symptoms of
myascheniagravisimprovc
224
11.1 lhe immunesystem I
~ mQmorycells
B-lyw(,ooc:)'leoow
+
cell divisionm
lormadoneol
activatedT-lymphocylebecomes
plasi tells cytoto~ic cell- destroy,
cellsinlectedbyviru,,
ororhercellscarry;ng
anti txxlyseaeted,
antigeom•,roome
ant~:;9~;~of
Figure 11.8 The roles of S- ar>dT-lymphcxytes;n the immull<' systl'ffi-a rnmma,y
Immune respo nse: the complex series of reactions of the lxxly to an antigen, such as a molecule
ontheoutsideofabi!Cterium,virus,parasite,allergenortumourcell.
• The immune response begim with an innate first response, carried out by phagcx:ytic white
bloodcells,whichcandestroyandengull(byphagocytosi,tendoc:ytosis}manydifferentfore1gn
organisms.
• At the same time, the primary phase of the adaptive immune system response begim, in
whichspecificck>nesofB-lymphocytesandT-lymphocytesdivideanddifferentiatetoform
antibody-o;ecreting plasma cells {from B-lymphocytes) and T helper cells and T killer cells {from
T-lymphocytes)thatarespecifictotheantigen,contributingtoitsdestructionorpreventingits
activity.
• This leads into the secondary phase of the adaptive immune system response, where memory
cellsretainthecapabilitytosecreteantibodiesoractasThelperorTkillercell:sassoon~the
specificantigenisdetectedagain
22S
m 1mmunity
Active and passive By the end of this section you should be able to:
immunisations are effective
waystotreatandprevent a) relatethe molecular,tructure ofantibodi~totheirfunction•
infectious diseases . b) outline the hybridoma method for th e production of monoclonal antibodi~
Smallpox has been c) outline the u,e ofmonoclonalantibodi~inthediagnosi,ofdi,elt<e andinthetreatme ntof
eradicated;otherdiseases
maysoonfollow,but d) di,tingui,hbetweenactive andpassive,naturalandartilicialimmunityand explainhow
vaccine development has
proved more difficult for
e) d iscus, the ,.,...ons why vaccination programm~ have eradicated smallpox. but not mea,le<,
diseases such as malaria .
tuberculosi,(TB),maluiaorchole ra
Monoclonal antibodies
We have seen that white blood cells in the blood provide our main defence against the invasion
of the body by harmful microorganisms. Special cells among the white blood cells, called
lymphocytes, are responsible for the immune response. Among these it is the n.ly mphocy1es
that secrete antibodies. Antibodies are very effective in the destrucrion of antigens within the body
(Figure 11.7). Mo n oclonal :uuil>odies are produced by a single clone ofB-lymphocytes . They are
identicalantibodieseffectivcagainstasingle, spedficantigen.
226
11 .lAntibodiesandvaccination .
Figure11.9Theanttxxly-a
quatema,yprotein
lymphoctye,areiK!ivated
______.. whenanant>genente«the ______..
bodyandbindsto arecepto,
onthecellsuffacemembrane
;nactM,
lymphocyte
andmaybeseaeted
byexocytosisa,
lree...tandir,gmok-cules
Figu"'11.10Compuler·
generatedimageofanan~body
molecule
227
m 1mmunity
4 Studytheproduction
of monoclonal
antibodies illustrated
in Figure 11.11.Make
aconciselistofthe !hesecel~.onc:eisciated, a,ecultufl'd
sequence of steps in andSUW'i'thoemonodonalantibody
their production
Figure11.11Theformationofmoooclooalantibodies
228
11 .lAntibodiesandvaccination .
AnO!her application of monoclonal antilxxlies in the tre:itment of disease is as ' passive vaccines '.
A monoclonal amilxxly specific to a pankular pathogen may be produced and injected imo a
patiem's blood circulation to good effect.
An issue to nOl:e is that, because the original monoclonal amibodies were developed from mice
cells (Figure 11.11), a patient may develop an adverse reaction to an antibody. This is because
the monoclonal antibody is itself a foreign protein to the patient"s immune system. Genetically
engineered antibodies that are compatible with the human immune system are sought to avoid
thetriggeringof theimmuneresponse.
abluecoloorawea15hefeilthe
urineninrajnsHCG(=pregnant)
immobileaoti bodlesuapthemobile
HCGaotibodH.><wi!hcolo<.xedgraoules.
soabl,,..roloorappears inwindow
coloufl'dg<anulesmo,eooupthetl'SI
mipandcombinewith!heimmobile
antibodieshere.gi,,;ngasecoodblue
coiou, coofirmingthe!estiscomple!ed
(rhi<roloor~,:,pea"wooheforoo r
rhereisHCGitJrheurine)
Figure11.12Detectingp,egriancyusir,gmonoclonalantibodteS
230
11 .lAntibodiesandvaccination .
231
m 1mmunity
Table 11.) Why vacc;nation ha, ool e,adic:ated meask>s, Til, mala,;a, sickle cell anaemia or cholera
lmpact(oroth.,.otherwise)of,..accination -andwhy
Thecholera-causingbacte,;umandilstwcinareactiveintheintestine - a,egklllofthebody"t,ere thevaccineadministeredbyinjeclion
cannot operate
The oral vaccine that ha, been developed offers pmtPCtion from some but oot all slfa ir,s of the cl,.ole,a bacter;um
{Standardhygiene prPCautions - deanwaterandefficientsewagedisposal - iftheycanbeachieved,arethemosteffectiveprotectklll
Cholera is endemic only in rnuntries where these rnnditions cannot be ma;nta ined .)
Noeffectivevacdnehasbeenproduced as yet. Thed iffirultiesare
• thepathogenisaprotoctist,withmanymo<egenesthanb~cteriaandviruses.Theallelesofthesegenes resultinvastnumbe"of
antigensbeingpresentonthesurfaceofthepathogen - indudingd ifferentantigensontheseparatestagesin thel ifecycle
• the pathogen spends a very short time in the blood plasma, before 'hidfr,g' from the immune system in liver or red blood cells in the
human host. This a ntige-n ic concea lme nt leaves little time la< an effective immune response
This is a genetic disease expressed in people wt.a are homozygous fa< the sickle cell haemoglobin allele (Hb')
Vacdnesardtheimmooeresponsec.annoteffectchangesintt...genetic:constitutioo(thegenome)
The effectiveness of the vaccine aga;mt tube,rulosis {BCG vacc;ne) varies accord ing to geography 80% ;n the UK, 14% ;n the USA
and;n,omeplacesitshowsnoprntectiveeffect
{Sttaimofthepatn<>genkbactedum,esistanttoanli·TBdrugshaveemergedinallrnuntries.)
Measles The disease persists because the virus is highly infectious and the vaccine is about 95% effectM', so a whole ;nfant population needs to
- a vaccine be treated with;n 8 months of birth ;n O!Uer to prevent ttansmission of the virus. Some children ,equ;,e one or ma<e 'booste~ injectkllls
exits befa<efull;mmunityisac:hleved,too
Many countries oow have about BO% of the population vaccinated but in African countries ii is rurn,ntly less than 50% that teeeive the
vaccine. For most communities, he rd immunity has oot been established
lnless..Jev<>lopedcoontrieswithahighbirth,atethelivingconditionsofmanypeoplea,ecrnwrled - idealconditionsfortransmission
of the virus. Al'iO, many victims a,e malnourished and the diets of infants are Jaw in protein {and deficient in vitamin N/. Any natural
resistancetoinfectionbythemeaslesvirusisminimal
232
Examination style questions I
• Defenceagaimtdiseasethatenterswhenthebodyis1\'0llnded stiITT.Jlateantibodyproductionandtheretentionofawrop-iate
andtheninvadedbyapathogenispro.idedby phagocytic memory cells
whitebloodcells.Theseareabletoengulfanddestroyforeign • Monoclonal an ti bod ies are a source of a single antibody
==
• The immunesystemisprovidedbyB-a ndT-l ymph ocytes
madebyaparticulartypeofB-lymphocytefusedwitha
cancercelltoforma hybridoma. Thisdividesindefmitely
which, when ,;emitised by foreign material (a nti gens), andsosecretestheantibodyinsignificantquantities
respond in complex ways to overcome the invasion, including Monoclonalantibodiesareusedinthetreatmentof
by the productionofspecifica nti bod iesthatdestroyor cancerbytargetingspecificda magedord iseased ce ll s
inactivate the foreign matter. MemOf)' cells are formed and andkillingthembythedeliveryofa'magicbullet'.They
retaintheabilitytorespondagainintheeventofre-infection are used in diagnmistoidentifythepresenceofspecific
e Vaccin atio n isthedeliberateadministrationofantigernthat macromolec ul es,asinpregnancytesting,andinseveral
havebeenreoderedharmlessbutneverthelessareableto other medical applications.
c:==::···
Fig.1.1
a)OnacopyofFig.1.1 :
i) Drawacirclearoundavariableregion [1]
ii) Draw in and label the position of the disulfide bonds
in the molecule [1J
iii)Explaintheimportanceofdisulfidebondsinprotein
molecules, such as antibodies [3J
b) Demibe how antibodies provide protection against
pathogens [4J
c) Other proteinsarefoundincellsurlacemembranes a) Nametheloiteoforiginofphagocyte1and
Demibethreerolesoftheproteinsincellsurface lymphocytes [1]
membranes 13) b) Name
(Total: 12) i) cellsA, BandC [3]
(Cambridge /ntemationa/ AS and A Level Biology 9700, Paper
ii ) organ D [1]
21 QI November 2009) c) Explaintherolesofthecells,A, BandCinanimmune
respome
2 a) Plasmacell1secretespecificantibodie1butare In your answer use the terms antigen and non-seff. (5]
short-livedinthebody. Explain,withuseofadiagram, (Total : 10]
how a plasma cell i:s made 'immortal' in the production of (Cambridge International AS and A Level Biology
a monoclonal antibody. [BJ 9700, Paper 21 Q6 November 2011)
b) Detailtheuseofmonoclonalantibodiesinone
applicationoftheiruseindiagnosis andoneapplication
inthetreatmentofdiseases. [BJ
{Total: 16]
233
A Level
~ 12.1 Energy
234
12.lEnergy .
- :'~,;o - ,,~-.0~..
t
energy transferred
toATP;nmany
r
ene,gylmtas small steps(ATP
"'" """°'"'
inmtJScu!ar
cootr;octioo)
- co,+ H,O
-- co,+ H,O
People sometimes mlk about 'burning up food" in respiration. In fact, linking respiration to burning
(combustion) is unhelpful. In combustion, energy in fuels is released, in a one-step reaction. as
heat. Such a violent change would be a disaster for lxxly tissues. In cell respiration. a large number
of small steps occur. Each is camlysed by a pa.n::icular enzyme. Because energy in respiration is
transferred in small quantities. much of the energy is made available to the cells. It may be trapped
in the energy currency molecule adcn01Slnc trlphos phatc (ATP) . However. some energy is still
lrnt as heat in each step (Figure 12.1). Jn animals that regulate their body temperature. this source
ofhe-atisimportant(Figure 12.13).
235
IEJ Energyandrespiration
• the active transport of molecules and ions across membranes, carried out by membrane pump,.
• movement by organisms. including those due to muscle contractions. Energy is also transferred
for movements within cells and by cell organelles.
• formation and secretion of substances from cells, including enzymes and honnone:s.
• cell d ivision. and the fonnation and maintenance of all the organelles of the cpoplasm.
1 lnFigure12.2the
structural formula
ofATPisshown .
lnFigure6.1
(page111)theway
the components
of nucleotides
are represented
diagrammatically is
shown. Use the symbols
inFigure6.1
to represent the adenosinecliphospha te AD?
structure ofATP. adenosinemooophosphateAM P
Figure12.2ATP.ADPandAMP
A1l' is referred to as "e n e r gy currcnq•" because, like money, it can be used in different contexts
and it is constantly recycled in 1he living cell. When the outennosi phosphate is removed,
adenosine diphosphate (ADP) is fonned and energy is transferred.
Later, it is energy transferred from respiration that refonns ATP from ADP and phosphalc (P,).
The reactions that yield energy and generate ATP from ADP and phosphate (PJ are particular sieps
inrespiration.Wewillreturntotheseshonly.
All these changes make up the ATP- ADP cycle (Figure 12.J).
The free e n ergy available in the step from ATP to ADP is approximately 30-34kJmol1. Some
of this energy is ]OS{ as heat in the reaction, but much of the free energy is made available to do
useful work - more than sufficient to drive a typical energy-requiring reaction of metabolism, for
example. Acrually, ATP may lose both of the outer phosphate groups, but nonnally only one is ]OS{.
It is only in a few reactions that more energy is required and in these cases both may be lOllt
An important way in which ATP serves as a reservoir of usable chemical e nergy is its role as
a o:immon ln1enncdia1e between energy-yidding re-acrions and energr-requiring re-acrions and
processes. We saw an example o f this in prolein synthesis (Figure 6.12, page 121). There are other
ways in which energy is transferred, including muscle contracrion (page 249).
236
12.lEnergy .
In s ummary. ATP is a molecule universal to all living things. It is the source of energy for chemical
change in cells, tissues 3nd organism.,. The impormm fe3rures of ATP are that it is
• 3 substance th3t moves e:isny within cells and organism., - by fad li1a1cd diffus io n
• formed in cellular respiration 3nd mkes pan in many re:iaions of mernbolism
• able to transfer energy in relatively small amounts, sufficient to drive individual reactions.
Jn the course d the A11'- ADP cyde in cells:
• All' is fanned from adeno,ine dipho,phate (ADP) and phosphate ions (Pi) by the transfer of
energy in respiration
2 OutlinewhyATPis
aneffic,entenergy • inthepresenceofspecificenzymes,All'isinvolvedinenergy-requiringreaaions
currency molecule We will return to the issue of just how the energy of ATP brings about useful work in cells and
organisms shor1ly. We will focus on the process of cellubr respiration first.
ADP+P;
~ I AT!'
metabolic:rei>clioosandmetaboHc:pf<xesses
driveobyeoergytraosfenedfrnmATP.e.g
• moYement of materials acr= c:ell membranes
• buildingmoleruH'S.indudir,gmac:romolec:uH'S
andotherbiomok-c:ules
• moYementsbymusclec:ontractkm.andother
c:ell mov,,ment,
Redox reactions take place in biological systems because of the presence of a compound
with a strong tendency to take electrons from another compound (an ox id is ing age nt) or the
presence of a compound with a strong tendency to donate electrons to another compound
(a r cd u d n g:igen t).
Anoiher feature of oxidation and reduaion is an e n ergy c h a n ge. When reduaion occurs,
energy is absorbed (it is an e n<lergon ic reaction ). When oxidmion occurs. energy is released
(an c x erg m lic reactio n ) . An ex:imple of energy rele:ise during oxidation i., the burning d
a fuel in air. Here, energy is given out :is heat. In faa, the amount of energy in a molecule
depends on its degr= d oxidation. An oxidised subsrnnce has less stored energy than a
redlK'ffi substance. An ex:imple d this is the fuel molecule meth3ne (CHJ which has more
stored chemical energy than carbon dioxide (CO,).
237
IEJ Energyandrespiration
' " ' ' ' · · - ' " ' " " ~ " " " ' 6-"___,,
" '°'"'
I
," ~actiooeod(hyd,ogen
A
,oo transport)
··'"'=''"' .•.,",,.,~
·"""'"""''''''~~
238
12.lEnergy .
3 Cocnzymc A (CoA)
CoA is involved in the removal of two-carbon fragment, from certain carbohydrates and lipids
during respiration, and also in food-storage reaaiOfls. You can see CoA '31 work' in the link
reacrionofcellrespimtion(Figure12.9).
·~~
~
;:::,
Figure12.SOthernudeobdederivativesinvolvedinrespirahoo
l
S-y
l'M>--Calbooffagmem
(ace~oott)lnuaospon
Lookuptheserefere"cesnow
Actually, the reactions of substrate-level phruphorylation occur twice in glycolysis, and it also
occurs in the Krebs cycle. However. the 3mount of ATP generated by this mechanism is trivfal.
2 The bulk of ATP synthesis mkes place in special scrucrures a.se-ocia1cd with the electron trnn..-..
port chain, present in the membrane-bound organelles, the mitochondria (page 18) (and also
ingreenpl3nt.sinthechloroplast.s (pages19-20), 3sweshallshortlydiscuss (see'Theelectron
transportchainandoxkbtivephosphorylation' , page246).
239
IEJ Energyandrespiration
Figure 12.6Acalorimete,lo,measu,ingenergyvalues
240
12.lEnergy .
Respiratory quotient
The respiratory quo1kn1 ( RQ) is the ratio of the amount of carbon dioxide produced to the
amount of oxygen taken in, in a given time, by an organism.
RQ =C~:=~
e.g.,foracarbohydrate:
Cv{,,06 + 60, --+ 6CO, + 6H,O
RQ = ~ = l
e.g.,foralipid:
2C,-,Huo06 + 1630, ....,. 114CO, + 1lOH,O
n4
RQ = - = 0.7
'"
Cakulationoftherespiratoryquocientisusefulbecauseitmayindirntewhichsubstratesarebeing
used. For example, when h exo;;esugarsarerespiredaerobically:
therespiraroryquO{iemis:
co,produced
RQ = o,rnkenin
= 616 = t
Where-as,whenfattyacidsarerespiredaerobically:
RQ =C~:=~
"'"
= - = 0.7
The respiratory quotient due to the respiration of fat (RQ = 0.7) is significantly lower than when
carod,ydratesarerespired(RQ = 1.0). Thisisbecausefatshaveagreaterproportionofhydrogen
relativetooxygen(theyareamorehighlyreducedformofrespirarorysubstratethansugars).More
oxygenisttquiredfortherespirationoffats.
The respiratory quotient when proteins and amino adds are respired is typirnlly 0.9. Where
proteins are used as the substrate, we have noted that the amino group (- NH :,) has first to be
removed (dcamlnation). In mammals the amino group is converted to urea and excreted. Plants
rarely respire proteins. If they d id , the ammonia gas formed would be hannful if not quickly Jost
fromthecellsoftheorganism.
De.spitethiscleardifferenceinvalue.sfordiffcrentrespiratorysubstrates,respiratoryquOl:iem
studies have only limited value. Rarely will an organism (such as genninating seeds) be respiring a
single respiratory substrate. Manyorgani.smsarefoundtohaveanrcspiratoryquOl:ientof0.8--0.9,
but this may be due to respiration of both hexose sugars (such as glucose) and fatty acids, in
varying proportions.
241
IEJ Energyandrespiration
4 Cakulate the re1piratory quotient of the saturated fatty i!Cid, palmitic acid {C ,.H ,, COOH)
5 The respiratory quotient {RQ) of two species of germinating seeds was meawred at two-day
intervals after germination
I Day 12 I• I• I• I
I Seedling A l o.65 l o.35 l o.48 l o.68 I
I SeedtingB l o.10 l o.91 l o.98 11.0 I
a Plot a graph to rompare the changes in respiratory quotient of the two specie1 during
early germination
b Suggestwhichrespiratorysubstratesmaybebeingrespired
12.2 Respiration
Respiration i1 the proms By the end of this section you should be able to:
whereby energy from
rnmplexorganicmolecules a) li<tthe four,tagesinurobkrespiration(9lycoly,is,linkreaction,Kre b<cy<l ea ndoxidative
istransferredtoATP. phosphorylation)and,tat e whe re eachoc:cur..ineukaryotk ce ll•
b)outline gly,:olysisasphosphorylationofglucose andthe ,ub<eque ntsplittingoffructose
1,6-bisphosphate (6C)intotwotriose phosphate molecul es,whkhue the nfurthe roxidi,edto
pyrwate withasmallyie ldofATP a ndre ducedNAD
c) explainthat,whe noxyge nisava ilable,pyrwate i,convertedintoace tyl(2C)coe nzymeAinthe
Aerobi c respiration
Cellular respiration which requires oxygen from the air is called aerobic n,splrn tlo n. Most
animals 3nd plants 3nd very many microorg3nisms respire aerobically. Tht,,y do so most, if not all
the time.
Jn aerobic respiration, sug:,r is o xid ised to carbon dioxide 3nd water 3nd much energy is
transferred. The steps o f 3erobic respiration are summarised by 3 single equation. This equ3tion
is equivalent to a balance sheet o f inputs (the raw materials) 3nd outputs (the products). II tdls us
notbingabou t tbe sleps.
242
12.2Respiration .
.
rH piratio n
, ::~
, ~o,,.,J ' \ :.': ': I ~·,;
\.. ~
-· t
------ ----·~~ --~;~;.._•
products 1 ca<bonckm,
-,~
Figure12.7Thertagesolaerobk,espirahoo
Glycolysis
Glycolysis is a linear series of reactions in which 6--carbon sugar is broken down to two molecules
..,...
of the }-carbon pyruvare ion (Figure 12.8). The enzymes of glycolysis are located in the cytoplasm
01Jtside organeUes (known as the cyro,;ol) rather than in the mitochondria. Glycolysis occurs in four
• Phospho ry la1ion by reactions with ATP. In this W3Y glucose is first activated. The product is a
s ix .carbon s ugar with two phos phate groups attached. NO(e th3t. 31 this stage of glycolysis.
rather than producing ATP. two molecules of ATP are consumed per molecule d glucose respired.
• l.ysis (splitting) of 6-carbon sugar phosph31e now takes pl3ee. forming two molecules of
three.carbon s ugar (called trirue phruphate).
• OxWation of the }-carbon sugar molecules occurs by removal of hydrogen. lbe enzyme for
this re-acrion (a dehydrogenase) works with 3 eoenzyme, nkotlnam.lde adenine dlnudeotid<'
(NAD). NAO is 3 molecule that can accepi hydrogen. It then becomes reduced NAD. (Luer.
reduced NAD will pass hydrogen ions and electrons on to other accepior molecules. When it
does, it is oxidised back to NAD)
• ATP ,-ymhesis. This happens twice in the reactions by which each triose phosphate molecule
is convened to pyruvate. This form d ATP synthesis is known 3S "at s ubstrate levd' . It is quite
different from the way the bulk of ATP synthesis occurs later in cell respiration (during the
operationoftheelearontransportchain, seepage246).
So. at this stage, as two molecules of triose phosprulte are converted to pyruvate. four molecules of
A1l' are synthesised. ln total. there is a net gain of two molecules of ATP during glycolysis
243
IEJ Energyandrespiration
~
1Phosphorylation
•·=
, ,, t:i
(uses AT!')
2Lysis
f:
~
(splittingol!he
6-carboobadbooe)
30x idation
+aalitioool
ioorgar,kphospna !e
J'=:~ll:t:::
:::~
:t~:~~a~:i~
~~u~~~Po:~=
~
:IIBEl
:~
='t::
4molecu~ofATP-aft'lormed
KK(g
=~:tilfinol2
Figure12.8Glyrnl;,,;is:as.ummary
As it is fonned during the llnk n:ae1 io n , pyruvare diffuses from the cytoool into the matrix of
a mitoc:hondrion. There it is metabolised. Firstly, carbon dioxide is removed from the J--carbon
6 Statewhichofthe pyruvate, a reaction called deearboxylation. At the same time, oxidation occurs by removal of
following are produced hydrogen, a dehydrogenation reaction. NAD is converted to reduced NAD during this reaaion.
duringglyrnlysis The product of the oxidative decarboxyla1ion reaction is an acetyl group (me-:, ning a 2-earbon
a CO, fragment). This a=tyl group is then combined with a eoenzyme called cocn7.yme A. forming
b lactate acetyl coe nzyme A.
c reducedNAO
d glycogen
e ATP
f NAO
g pyruvate
h glucose
The production of acetyl coenzyme A from pyruvare is known as the link reaction because it
connects glycolysis to the reactions of the Kreb. cycl e. which now follow.
In the Krebs cycle (Figure 12.9), the acetyl coenzyme A enters by reacting with a 4-carbon
organic add (oxaloacetate). lbe products of this reaction are a 6-carbon add (citrate) and, of
course, coenzyme A. This latter, on release, is reused in the link re-:,aion.
The Krebs Cycle is named afrer H ans Krebs who led the team that discovered it. It is also
sometimes referred to as the citric acid cycle. after the first imennediate add formed.
244
12.2Respiration .
Figure12.9Thelinkreactionandlvebscyc:lecasumma,y
Next. the citrate is convened back to the 4-cubon acid (an accepior molecule, in effect) by a series
of reactions. This is why Krebs is called a 'cycle'. These are the changes.
• Two molecules of carbon dioxide are given off. in separate dernrboxylation re-actions .
• A molecule of ATP is formed, as pan of one of the reactions of the cycle. As is the case with
glycolysis, thisATP synthesis is at substrate level.
• Three molecules of reduced NAD are formed in dehydrogenation reactions
• One molecule of another hydrogen accepter, FAO (flavin adenine dinudeotide). is reduced.
(NAO is the chief hydrogen-carrying coenzyme of respiration but FAD is another coenzyme with
thisroleintheKrebscycle.)
Finally, remember that glucose was convened to two molecules of pyruvate in the glycolysis phase.
This means that the whole Krebs cycle sequence of re-actions 'turns" twice for every molecule of
7 Outlinethetypesof
reactionCiltalysedby glucosethatismetabolisedbyaerobicrespiration.
a dehydrogenases We are now in a position ro summarise the changes to the molecule of glucose that occurred
b decarboxyla1es in the reactions of glycolysis, the link reaction and the Krebs cycle. The product.s formed during
glycolysisandtwoturnsoftheKrebscycleareshowninTable12.I.
Table12.1Netproductsofaerobic:respirationpermoleruleofgllJCoseattheendoftheK,ebscyde
245
IEJ Energyandrespiration
You can see that, at most of these events, NAO is convened to reduced NAO. However, at
one point in the Krebs cycle. an alternative hydrogen-acceptor coenzyme (known as FAD) is
reduced.
Now. in the final stage of aerobic respiration, the hydrogen electrons are transported along a
series o f carricni. from the reduced NAO (or FAD) produced in the e:irlier stages. The terminal
acceptor is oxygen. Now. finally, hydrogen has been oxidised and oxygen reduced to water
(Figure12.10).
The 'carriers' (they are conjugated proteins - page SO) are located in the inner membrane
of the mitochondria. These carriers occur in highly ordered sequences. each consisting of
hydrogen acceptors and electron carriers next to a final enzyme, ATP synth asc (known
for short as ATPase). The location and physical structure of these systems are illustrated in
Figurel2.ll.
Theoretically at least. the coml yield from aerobic respiration is about 38 molecules of ATP
per molecule of glucose respired. In pmctice, it is less than that. probably about 32 molecules
ofATP.
8 Distinguish between the
following pairs
a MJbstrateand
intermediate
b glycolysisandthe
Krebs cycle
-jz·· · · · · · · · · · · ·~ -
t oxidationand
reduction
;E;F
Kreb<c)de
\
=::~=lrom '7 \ ener!i=~f:•d
.) =--~-- ~~9~~1:·
wheo,educ:edFADisoxi~. 2.5ADP+3P;
make All' ;:;r~~-
is caned
ooly1.5moleculesofATParefomll'd metabo lic water
246
12.2TheprocessofATPsynthesis I
4' 12.2 continued... The process of ATP
synthesis
247
IEJ Energyandrespiration
--
::::=~ &::=z:s:::::JII-
..:. ,·~~~
! ATPase
ATPsyntr.e<isa,upledwithpromn
flowdowllcOflCeotra1""1gradieflt
This process by which ATP is generated has become known as c hcmi01>m 01<i~. It was a
biochemist, Peter Mitchell. who suggested the chemiosmOl:ic theory. This was in 1961. His
hypoihesis was not generally accepied for some years but, about two decades later. he was
awarded a Nobel Prize for his discovery. Today there is a great deal of evidence for the occurrence
of chemi0.5mosis. We can describe it as a correct hypoihesis. So the word 'theory" in its title is used
initsbroadestsense - asanacceptedo:incept.
248
12.2TheprocessofATPsynthesis I
Case studies of ATP 'at work'
I An example of an anabolic reaction of metabolism 'driven' by ATP is the manufacture of
pr{){eins in the ribosomes. Each amino add used to build up the primary structure of the
procein is first atrnched to a specific molecule of transfer RNA. ATP is required for this
'activation'reaction(Figure 6.12,pagel21).
2 ATP also reacts directly with protein pumps in membranes, so providing the energy needed ro
changetheshapeoractivatetheproceinandsomoveionsand01hermetabolitesacrosscell
membranes which would olh.erwise block their movement. The movement of a single substance
is iUustrnted in Figure 4.21 (page 91). 1he sodium----potassium pump is shown in Figure 4.22.
j Sometimes ATP reacts with water (a hydrolysis reaction) and is converted to ADP and a
phosphate ion. For example, direct hydrolysis o f the terminal phosphate groups like this
happen in mu..-.clc contniction.
Muscles that cause [ocomo{ion, skdeml muscle, consist of bundles of muscle fibres. 11iese fibres
are long multinudeate cells that are able to shorten to half or even a third of the relaxed or
'resting' length. lbe whole muscle fibre has a 'stripy' appearance.
This is due roan interlocking arrangement of two types of protein filaments, made of the
pro1eins my01<ln and actin.
111ruonmlcrographolLSthroughpartola
ok1l1tal muscl1 llbr9. HP(xl0000)
n
change In a slngl1 sartomer• In r<1land and contucwd myollbrN
M<lsdescom,aaas!heacllnaod"'JOS#lfllilmmtssllde
betweeneachothef.soortffl119eachsarromefl'
(Continued)
249
IEJ Energyandrespiration
(Continued)
Look at Figo,re 12.12 now so that)"" cm, understand "'here ATPoperates to bn·ng about muscle
SkeletalmusclecontractswhentheactinandmyosinfibmentsslidepasteachO(herina
series of steps. sometimes described as a 'ratchet mechanism'. A great deal of A11' is used
in the contraction process. This cycle is repeated many times per second, with thousands
of myosin heads working along each myofibril. ATP is used up rapidly and the muscle may
shorten by about 50percentofits relaxed length .
4 Another significant application d the energy transferred in respiration concerns body
temperature. The major so1.m:e of heat in mammals (and in birds - both groups o f vertebrate
which regulate their body temperature) is as a waste product of the biochemical reactions of
metabolism. Heat is then d istributed by the blood circubtion.
The organs of the body ~"'Y greatly in the amount d heat they yield. For example, the
liver is extremely active metabolically, but most o f its metabolic reactions require an input
ofenergy(theyareendergonicreactions)andlinleencrgyislostasheat. Mostly,theliveris
thermally neutral.
When at rest, the bulk of our body heat comes from cxher abdominal organs, mainly from
the heart and kidneys, but also from the lungs and brain (which, like a computer central
processingunit,needstobekepicool). Underrestingconditions,theskeleton,musclesand
skin, which make up over 90 per cent of the body mass, produce less than JO per cent of the
body heat (Figure 12.13). Of course, in times of intense physical activity, the skeletal muscles
generateagreatdealofheatasawasteproductdre-spirationandcontraction.
organ ma .. heat
(as % ofthe production
w""""'
=>)
atrest(as %
of !he total)
10 \/v'hatdoe5itmean
when we s.ay most
reactionsinlivercells
areendergonic.
Fig..-e12.13Heatproductioointhebodyat,est
25-0
12.2TheprocessofATPsynthesis I
The site of cell respiration - structure in relation to function
The location of the stages of cellular respiration are summarised in Figure 12.14. You can sec, that,
once pyruvate has been formed from glucose in the cytosol, the remainder of the pathway of
aerobic respiration is located in the mitochondria. lne relationship between strucrnre and funcrion
inthisorganelleissununarisedinTable 12.2
!hel'flzymesoithe linkreactioo
:;;!~~1,:;;:ebs()dearelocated;o
theen,.,.,...,olglycolysis - · · ADP
a,eloca1ed;n1rn,cy1oso1 +P;
glycolysis
glucose - - -~ - ~ - pyruvate- -
reduced
'" s.ilesofoxidatioool
hyclrogeoMCep!""
(waterformatk>n)
outermemb<ane andATPsyn!hesis
im,ermemb<ane
Figure12.14Thelocahonofthe,ugesofcellular,espiratioo
extemal double membrane permeab~ to pyruvate, CO,. 0,, NAD and redtKed NAO
siteoftheenzymesofthelinkreactionandtheKiebsc)'Cle
• surfac:eareagreatlyincreasedbyintudingtolormcristae - sincetheelect,oo
transportchainandATPsynthaseenzymesarehoosedhere,opportunit>esfor
ATPsynthesisareenhanc:ed
• impe,meabletohydrogeniom{protons) - allowingtheformationofa
potential difference between the inler·memb,ane space and the matrix
inte,-memb ranespace 1elativelytinyspace - a lkM'ingtheaccumulabonofhyclmgenKlm(protons)
tfie1etog.eneratealall_ll'polentialdifferencewiththematrix,making
phmphmylationJ>=ible
2SI
IEJ Energyandrespiration
12 0etailwhatchanges waterbathsinarangeoftemperatures.
are required to this egat202S303540and50°C
experiment in order
to investigate the
effectof§Ubstrate "''
--00
concentration on the
aclivey,,ast
rate of respiration
wrthmethylene
~ bluesolutoon
Figu~12.15Aninvestigationoftherateof,espiratiooinyeast
theamoontandcoocentrationofthe)'<'astsuspension
thevolumeofoilandmethylenebluesollrtklo
independ e ntvariabl e therangeoftemperatu,es
dependent variable thetimetakenfordecoloufisatkloofthemethylene
bi~
252
12.2Anaerobicrespiration .
Anaerobic respiration
Many organisms are able ro respire in the absence d oxygen. at least for a .short time. 1bcy do so
by a process ralled fermentation or anaerobic respiration. Some tissues in an organism may respire
anaerobirally, too, when deprived of sufficient oxygen (see below).
In the absence o f oxygen, flowering plants may respire by akoholk fermematlo n , for
example, the cells of roots in waterlogged soil. Many species of yeast (Saccharomyces) respire
anaerobirally (some do so even in the presence d oxygen). The products are ethanol and rarbon
dioxide. Akohollc fenne ntatio n of yeast has been =ploited by humans in wine and beer
production for many thousands of years.
Once again, these equations are balance s h et.'ls of the inputs (raw materials) and the output
(products) o f cellular respiration. Tht,>y tell us nothing about the separate steps involved, each
catalysed by a specific enzyme. lbey do not mention pyruvate, for example.
2Sl
IEJ Energyandrespiration
Figure12 .16Therespiratory 2x lx
pathways of anaerobic glua>H- r y !hrne-carton: : z r - : p)'uvate ••••••••• 6C0,+6H20 (aerobic
fl'Spiraticm
sugar 2xNII.D ,,;.,,;o,
pathway)
ni;t;:r_;t rerucedNAD
lxe11,.anaj2xc0:,
Here<tt ha na l tl!he Herepyruvateis !he
s~:~
acceptor for reduced redocedN/l.0 --..... / ...-- ,educ:edNAD accep1ortor ,edoced
NAD alcohol dehydrogenase lactate dehydrogenase NAD
2xN: z
A few organisms re.spire only in the absence of oxygen. One is the tetanus bacillus CI-Ostridi,m,
14 Canyouruggest /eta11i which causes the disease lockjaw. This bacterium is active in oxrgen-free environments
a mechanism by rich in nutrients. These conditions occur in deep body wounds. Otherwise the bacrerium
which the absence of
merely survives as resistant spores in the soil or among the faeces d the lower gut d many
oxygen1nre1pmng
animals. During growth (but nOI: as a dormant 'spore'), C. telm,i produces toxins which, when
tissuemight'switch
off'boththeKrebs released inside wounds, are carried around the body in the blood circulation, causes agonising
cycle and oxidative muscular spasms. Today. tetanus can be prevented by vaccination, supported by periodic
phosphorylation? booster injections
254
12.2Anaerobicrespiration .
Glycolysis
(net production at the (re-oxidised (net production at the
substrate level) during substrate level)
lennentabon)
(producedbyoxidativ,,
phospho,ylation)
(produced by oxidative
phosphotylation)
Krebsc)'CH'
'(net production at the
substrate level)
(producedbyoxidativ,,
phospho,ylation)
(producedbyoxidativ,,
phosphoty\ation)
M3ny species of yeasc cannot metabolise ethanol; the energy locked up in et:l,3nol rem:,iru
un3vaibble ro the cell. This m:,kes these species indU51rfally useful organisms in the producrion of
ethanol. However. oiher yeast species can met3bolise et:l,3nol when cxher respiratory substrates are
used up.
2SS
IEJ Energyandrespiration
Secondly, the roots can and do respire anaerobirnlly by alcoholic fermentation, when =='Y·
The ethanol produced in the cells may be metabolised by enzymes present, but the root cells of the
riceplamhaveahighertolerancethanothercerealsforethanol.
paddyfieldcultureofrice - planringourofriceseedlings
15 listthefeaturesofthe
riceplantthateoable
itto,urviveswamp
conditions.Annotate
yourlist.~ thatthe
value of the adaptation
is made dear.
Figure12.17ThestrtKtureofmature,ice~ants.as""9etat>Yegrowthgi>eswaytofruitformation
256
12.2Anaerobicrespiration .
2S7
IEJ Energyandrespiration
Thel"!ukldropmoVl'Sdue
toalallinpres:s.ure;isthe
\Olume,Sreduc:ed,e.9
'Mll'JICO,gi,,..noot,S
atisorbedbythes.oda
lime.ar>do,istakl'Jlup
by the seeds
8\/lachan.ge
intemperature0<
atmospheric pressure
mi!jhttlavethesame
effect.andMl\dingol!he
appa<.rtusmayaddheat
andchangethere-.><!ing,
4 :::;/~n~":1~t:~~.~':.ed
maoometer.andthe"""-meolO,
16 lntherespimmeter absorbed,Sre.-!offonthesyfing,e
~ninFigure12.20,
explainhowchan~
graduatedsj<inge
in temperature
or pressure in the
external environment
are prevented from
interfering with the
measurement of
oxygenuptakebythe
resp1nngorgarns.ms.m
the apparatus cooncitube
(thermobarnmeter)
1 therl'5piriog orgaoi<ms
glassbeads(s.ame g1,,.. 011co,aod
volume;iss..>mplein absorb 0,
rP<P,mmetertube)
2
AP.ycllar\geinpres,;,.xeor temperaturedufing U-tubemanometer ~~~bedby
theexperimentallerutheresp<rometerrube 3manometerflu.klmoYP<totheright
andthethermobarometer tube equal~ due to the fal ,nprP<sure"'
thevok.mei<reducedbythe
byuptakeoloxyqen
Figure12.20Adiffe,ential,PSpirnmeter
258
12.2Anaerobicrespiration .
between1 0a nd 35°C above 40°c lbe differential respirometer is used with a water bath. By
thera1eofresp;ra1iooa1 thera!eol respi<alioodecreases
repeating readings at a rangeofdifferenttemperarures,theeffect
leastdooble,;forew,y withariseinll'mperature,
::;c::.:~7,:Jt: owingto<lena!Yfa!ioooi!he
enz;mesa1 hgie,1empe,atu!l'<
oftemperatureonrespirationcanbemeasured. Data such as that
inFigure12.21canthenbeobrninedwiththisequipment
Why does temperal!<reaffect the rate of respiration?
~
The enzpnes of respiratiOfl are proceins. All proteins are
denaturedbyhighertemperatures,althoughthetemperarure
atwhichproceiruaredenaturedvarieswidely. For=ample,
somebacrerialive successfullyinhotspringswherethewater
temperature may approach that of boiling water.
From the graph in Figure 1221 we can work out the
temperature at which significant denaturing of respiratory enzymes
in the organisms used in the experimem takes place. Below this, a
rise in temperature causes the rate of respiration to increase. It is
a feature o f chemical reactions that their rate at least doubles for
20 JO
every10°Criseintemperature.
tempe,ature/"C
Figu n, 12.21 Theeffectoftempe,atureontherateofrespirabon
259
IEJ Energyandrespiration
<D'"' J
plants and animals
,f)'N ~N B [Total: 15]
(Cambridge lntemational AS and A Level Biology 9700,
Paper 04 Q9 June 2008)
~rn, 4 Outline and explain
~o~
a) the difference between substrate level phosphorylation
andoxidativephosphorylation,
b) whyaerobicrespirationisdescribeda1moreefficientthan
!H OH OH
anaerobic respiration,
c) how reduced NAO is reoxidised in anaerobic respiration
5 Duringtheprocessofglycolysis,glucoseisconvertedbya
seriesofstepsintot1wmoleculesofpyruvate
~ Thediagrambelowoutlinesglycolysis
Fig.1.1 gluc1,::)
1
a) i) Namethenitmgenousbaselabelled B [1]
hexosephospha1e(6C)
ii) NamethesugarlabelledS. [1]
b) ATPisdesnibedashavingauniversalroleastheenergy
currencyinalllivingorganisms.Explainwhyitis
J""'
described in this way.
c) State precisely twopli!CeswhereATPissynthe!isedin
cells
[4]
[2]
[Total: 8]
/'~
(Cambridge International AS illld A Level Biology 9700,
Paper04 Q6 November 2008)
--r:': step 4
"""-r:"':
reduced NAO redocedNAO
2 a) Forthestagesofaerobicrespirationlistedbelow,identify
the1iteoftheprocessandthediiefproducts imermedir:'ate(JC:AOP+ 2 Pi intermedi~C:AOP+ 2 Pi
ofei!Chstep.
steps
2ATP J"-2ATP
Component
of process pyrwate(JC) pynNate(3C)
of aerobic process in the
respiration eukaryoticcell O.iefproducts a)Withreferencetothediagram above,statetheprocess
glycolysis occurring at
i) stepsland2 [1]
ii)step3 [1]
Krebscyde iii)step4 [1]
b)Explainwhygluco1eneedstobeconvertedtohexo1e
bisphosphate [2]
c) Pyruvatec.anenteramitochondrionwhenoxygenis
[12]
present.
b) \Nhatdoyouunderstandby
Describewhathappenstopyruvateinayeastcellwhen
i) decarboxylation [2[ oxygen is not present [4]
ii) dehydrogenation [2[ [Total: 9]
[Total: 16] (Cambridge International AS and A Level Biology 9700,
Paper41 Q6 November 201 I)
260
A Level
1 3 Photosynthesis
Photosy nthesis is th eene rgy tra nsferpr0<ess th atis th ebas isof Stud e nts use the irknowle dgeofpla nt ce ll sa nd leaf structure from
much o f life on Ea rth. It provides the bas is o f most food cha in s th e topi c o n Ce ll stru ctu re w hile study ing photosynth esis. Vario us
providing e ne rgy di rectlyorindi rect ly for a ll o the rorga ni sms. e nv iron menta l facto rs influe nce the rateat w hic hpho tosy nthesis
ln e ukaryotes, the processo« ursw ithin chlo ropl ast s.
261
IEJ Photosynthesis
to energy in the chemical bonds of a range of organic molecules. including sugars and ATP.
We summarise photosynthesis in single equations
1 What do we mean carbon dioxide + water + energy --+ glucose + oxygen
by'anabolism'and 6Co, + 6H,O + energy --+ c,,tt,,~ + 60,
'catabolism'7
Compt;re this eqo,a/ion with that for aerobic ,·espirm ion (page 243).
At first sight it seems that the one process is simply the reverse of the other. In fact these equations
are merely 'balance sheets' of the inputs and outputs. Photosynthesis and respiration occur by
different pathways with many enzymes unique to one or the other pathway. And of course,
photosynthesisis:inanabolicprocess,whereasrespirationisacatabolicone
lbe sugars produced in photo.synthesis provide the carbon 'skeletons' for the spnhesis of aU the
mctaholim• the plam needs . We can say that green plants are 'self-feffling" (amotrophic nutrition).
This type of nutrition contrasts with the ways animals and other organisms get their nutrients. In fact,
it is green plant nutrition that suppons the living world as a whole. Green plants are the source of
nutrients for almost all other organisms, d irectly or indirecdy. Jn this topic we look at photosynthesis
asancnerg}lransfrproa, s.
262
13.1 Photosynthesisasanenergytransferprocess I
Investigating chlorophyll
Scme plant pigments are soluble in water but chlorophyll is not. Whilst we cannot extraa chlorophyll
fromleaveswithwater,itcanbe=tractedbydissolvinginanorganicsolventlikepropanone
(acetofle). Withthissolvent,extractiondchlorophyllisfairlyscraightforward(FigurelJ.2)
2 ~"""'ffrlelyru!and 3aboot20onlolpmpanooe
placedinamona; (acemne)adiedassolveo~
!heogrour.dupwilh andtheconlel\tsground"'}ilin
asmallquantityofwashed !Oproduceaoonc:entrated
anddriedsa»d(asabra,Ne) <h~lextrad.
measuringc-,iiJ>de,
-t~=a--~,,~:_
Stubescentrifugedro 6extractedpjgmemsolutioocombined
(emoYesuspendedorganic andstoredintr.erold,io!hedar'<,
4 solution decamed into !WOCl'fltriluge tubes aOO matte< (w~l lragments, aschk>roph~I, are unstable molecules
~
Figure 13.2 Stepsintheextractionofplantpigments
263
IEJ Photosynthesis
As the solvent from moves up the chromatogram. components of a mixture move at different rates.
This is because they have different solubUities in the solvent. Also, the components of the mixture
beingseparatedinteractwiththematerialofthestationaryphasetodifferingdegrees.
"'"
Before the solvem front reaches the opposite end of the chromatogram, the process is stopped.
The level reached by the solvent front is marked. The chromatogram is then dried . The positions of
coloured components of a mixture are easily located (as in the case of chlorophyll). The locations
of any components that are nOI coloured have to be identified: various techniques are available.
p.aperchmmatog,aphy bu»g
ru,.
t ····..
(averyquickseparation)
Using:
~:
Thin.layer chromatography
dilor~~I
• lhin-la)'erchromatcg-aphy'J>late'
(pla<ticfilm)pre-coa!edwith
silkag•I
o chromatography'tank' -llat-
bottomedtubewithdose-fittiog
lnd..-,tifyingpigm.,,,tsbyR 1va lu..
Thedisranc:eaparticularsubstaoce
=..:,~~l~~~:;:'t·~:·
val,,.isknowoa:stheR,vah.e
R,= d::i'.:'::emoYedd\"'!e"..,7
chromatogram
···-··· - solven!frool,
=•11:.::i.y
runwa,,top!)l'd
loading spot
Figure13.3Preparir,gandrunn;regachromatogram
Look at the chromatogram in Figure IJ.J. Green leaves contain a mixture of phoiosynthetic
pigments. These include two types of chlorophyll, known a, c hlorophy ll a and c hlorophy ll b.
The others pigments belOflg to a group of compounds called caro tcn o lds . Incidentally, larger
samples of these individual pigments can be obtained by column chromatography. for =ample.
Such samples may be large enough ro investigate the propenies of individual pigment m olecules.
1be chemistry of the chlorophylls is pankularly important because they are directly involved in
the energy transfer processes in the chloroplasts. The stru<1ure of chlorophyll is .shown in figure 13.4
Looka11bestrue1ureofcb/oroph;dlnow.
The other pigments. the carotenoids, are described as accc ssor)· p igm e n ts . This is bemuse they
havetheroleofpassingthelightenergytheyabsorbontochlorophyll. beforeitrnnbeinvolvedin
phOlosynthesis. (One of the caroteniods is a yellow pigment called carotene - an important source
of vitamin A in human diets.
Thechlorophyllpigmentsoccurinthegrana,heldinthelipidsandprotein.softhethylakoid
membranes. Chlorophyll is large - a 'head and tail" molecule. in fact. The 'heads' are hydrophilic
and associate with proteins in the thylakoid membrane. The 'tails' are hydrophobic and associate
with lipids in the thylakoid membranes. Also found here with these pigments are the enzymes and
electron-carrier molecules involved in the steps of photosynthesis where light energy becomes
chemical energy. We will meet these shortly.
264
13.1 Photosynthesisasanenergytransferprocess I
chlorophyllsaandb lnthechk,,op!ast.chlorophyllsaandb
Thisischlornphylla: occur inthemembranesolthegrana - they
chk,rnphyllbhasan aresandwkhedbetweentheproteinand
Cl-13 aldehydegroup(-CHO)
;ntheplaceofthe :r._~}> l,ed I~= ofrhe =mb,T
- - headocru"
associated
withproteim
res.t1~sshowthat the
wa,,..ler,gthsofl>ghtabsomed
b) actionspectrum bjpholosynthe!icP".Jmem,
a)aboorptionspectrum ,ecordofamoum (largelyredaodl,jue)i>l"e
meas...-edllSOr,gaspearometer olphotosymhesisocrnn;ng very,;mila,tothewavelength,
a1eachwa""lmgth !llatdriYepho!os)'J\lhesi,
!
f
•
I
tili.o,light ,edllg,t
Figu re13 .5Absorptionandactionspectraofd,lorophyllpigments
265
IEJ Photosynthesis
The rate of photosynthesis at diff=t wavelengths may then be plotted ot1 a graph on the same
scalea.stheabsorpi:ionspectrum(Figure1J.5b).Wehaveseenthatblueandredlightaremost
strongly absorbed by chlorophyll. From the action spectrum we see that it is these wave lengths
thatgivethehighestratesofplxxo.synthesis
lig ht e ne rgy
Photosynthesis in chloroplasts
Figure13.6Thetwostagesofphotosynthesis:asummary
266
13.1 Photosynthesisasanenergytransferprocess I
At the same time that reduced
(Nlwtlnamldeadenln• dlnu<ltotlde phosphate) NADP is formed in the
light-<lependemstage,ATP
isgeneratedfrom ADP and
phosphate. Light energy is
n=dedforthis,too,ofcourse.
This energy comes from
photoacrivatedchlorophyll,
andtheprocessisknownas
pho to phosph<>r)·lalio n .
Also, o xygenisgivenoffasa
wasteproducrofthelight-
dependentsmge.
This stage of
photosynthesis occurs in the
gr:maofthechloropbsis .
scructuresinvolved (the
photOiSys{ems)next,and
thenatthestepsbywhichit
The photosystems
O,Jorophyll molecules do
nor occur haphazardly in
thegrana.Rather,theyare
groupedtogetherinsirucrures
Figure 13. 7 NADP and NAD called pho to,,.~·,ncm s, held in
the thylako id membranes of
light absorbed
the gra na (Figurel3.8).
ln e achphO(osySlem,
several hundred chlorophyll
molecules plus accessory
pigments (carotenoids) are
arranged , grouped together.
All these pigment molecules
many pigment molecules harvestlightenergy.andthey
{chlomphyllar>da<:cesso,y funneltheenergyto a single
pigments) chlorophyll molecule of the
photosystem. This molecule
is known as the reac ilo n
centre . The different pigments
aroundthereactioncentre-s
reactkmcentreof absorblightenergyofslightly
chlorophyll a differentwavelengths,butall
(abso<bir,gene,gy 'feedinto' the re-acrioncentre.
at700nm - PS1,
Nowwealsoneedtonote
orat680nm - PSII)
thatthere aretwo typesof
photosystempresentinthe
exited electrons thylakoidmembranesofthe
releasedffomhere grana . These differ in the
and replaced by
lowene,gy wavelengthoflightthatthe
{•ground-state) chlorophyll of the reaction
electrnffi
Figure13.8Thestructu,eofaphotosystem
267
IEJ Photosynthesis
• Pho tosys1c m I (PSI) has a reaction centre activated by light of wavelength 700nm. This reaction
centreis3lsorefcrredroasP700
• Pho t01Systcm II (PSIOhas a reaction centre activated by light of wavelength 68onm. This
reactioncentreisalsoreferredtoasP68o.
Photosystems I and II have differing roles, as we shall see shortly. However. they occur grouped
together in the thylakoid membranes of the grana, along with specific proteins, many of which
function as eke1ron .c:,rrkr molecules.
l lbe light energy reaching the re3ction centred photosystem [I activates a panicular pair of
ground..,.1ateclcctronsinthespecialchlorophyllmolcculethere. Theseareraisedtoan
excited sta te and immediately captured by an electron accqxor in the nearby chain of
electron-carrier molecules (Figure 13.9). By the loss d electrons the chlorophyll molecule is
oxidised, causing it to inunedimely acquire electrons from water. h is electrons in the ground
state that fill the gap in the chlorophyll molecule vacated by the =cited electrons. By this
process, the water molecule is split. The remainder d the water molecule is released as ox}"gen
gas (two oxrgen atoms combine to form a molecule of oxygen) and h ydroge n ions.
Meanwhile, the =cited electrons are passed along a chain of electron carriers. As this
happens, the energr level of the =cited electrons progressively falls back to the ground state.
Hmvever. the energy transferred from them as they flow along causes the pumping of hydrogen
ions (protons) from the chloroplast's stroma into the thylakoid spaces. Here the hydrogen ions
accumulate - incidentally. causing the pH to drop. The result is the creation of a proton gradient
acro.ss the thylakoid membrane - of which more shortly.
light energy reaches the reaction centre of photosystem I at the same time. Here, light
energy again causes the tjection d panicular ground-state electrons, raising them to an =cited
state. These , roo, arecapruredbyanelectronacceptorinthenearbych3indelectron-carrier
molecules. Now there comes a difference.
The gap vacated by these electrons in photosystem [ is filled by electrons that have been
transferred from photosystem II. These have now fallen back to the ground state energy level
~~~~~ -- - used!OdrlveATI'
synthesls(see
flgurell.10)
Figure13.9Zdiagram,oo,,,,,;ng
the light-dependent stage of
photosynthesis - - - 1"""'9"o!electroo,1romwa!eftol,IMJI' - - - - ~
268
13.1 Photosynthesisasanenergytransferprocess I
Meanwhile, the =cited electrons from phowsystem I are passed along a short chain of electron
carrier molecules and their energy is transferred to fom, a molecule of reduced NADP from
NADP and hydrogen ions, in the Sll'OOl3. Jn Figure 13.10 you can see that this occurs on the
stromal side of the thybkoid membrane - an important point.
P
phot<>S}ffi'ffi I
Re~cedNADPandATPa,e,equi,edforCO,.fixationin
thestroma .Throucj> prnductsofthehghtreactDn(taking
pla<:e tnthethylakoidmembrane),theyareactuallylormed
onthestromalsicle,wheretheya re ,eq u,red
.,:
ADP +
//Wt'>-. "-
y ATP
ATPsynthase
(ATPase)
W r.:::~::m . . 1
Figure 13.10Theorganisationofthethylakoidmembrane
269
IEJ Photosynthesis
chloroplast ::io~
- - ~ - - - - - - - -- ,educed NADP
NW+P~
\'I
\
'
~'-"'"'
rubisro"' fixation
·µ~
glycerate
/
C--C-C+C--C--C
2 x 3.carbof\
compoornl
calvin
cyc le
'"""~-,w:, ADP+P
.
'
o-ansloc:atedtorP<t
of plant
Figure 13.11The<eiKOOr6oflf>elighl~ndependentslageinsiw
270
13.1 Photosynthesisasanenergytransferprocess I
In Step lbrec, r q :c n cr:ulo n o f 1.hc acrcplor m o lec ule occurs. For every six molecules of trio.re
phosphate formed, five are used to fonn three molecules of RuBP. In this conversion, energy
from All' is also required. From the O(her molecule of triose phosphate Olher compounds are
synthesized. These include carbo h }"d ratc~ (sugars and starch, and sucrose for translocation in
the phloem), lipid~, and a mino acids. In these conversions energy from ATP is also required.
How the products of photosynthesis sustain the whole mctaholls m of the plant is summarised in
Figurel3.l2
------/
\ \
carboodiadde io ai,
""
k,,;sofwatef
.1orna,ealsoca<ried!othe""""'wheremuc:hamoooacid
symhesisocru"
•Mar'fvil.lmiosaresynthesi<l'dinthe<hloroplasts100
Figure13.12Photosynthesis:itsplaceinplantnutrition
271
IEJ Photosynthesis
Figure 13.13Acompafisonofcyclkandnon-cydicphotophospho,ylabon
-"°
a innon-cycl ic
plxxosymhesis in just the same way as unlabelled carbon dioxide ( ''co,) is. It was then possible
phoro-
to discover the sequence of metabolites which become labelled - in effect, the path o f carbon
inphomsynthcsls,frorncarbondioxidetosugarsandthea1herproductsofpho1nsynthesis
b incydicphoto-
A culture of Cblorella, a uniceUular alga, was used in these experiments , in place of mesophyll
phosphorylation7
cells. This was because they have similar photosynthesis and they aUow easy sampling. Samples
9 Ina school laboratory
ofthephotosynthesisingcells,takenatfrequentintervalsafterapulseof"CO,hadbeengiven
demonstration of
were harvested and analrsed. lbe intermediates that became progres.sivcly labelled with the
theHillrei!ction
{page277},what carbon--14wereisolatedbychrornatographyandthenidentifiecl.
electron acceptori,
used,andwheredo
the electrons come
from7 Chemosynthesis
10 Manybiologist,are
Autotrophic organisms are 'self-feeding' organism.,. lbey fall into two distinct groups, according
not surprised the
enzymerubiscoi, tothesourceofenergyusedtobuildcarbondioxide into carbohydrates
the most abundant One group, the photosynthetic organisms, use light energy. We have focused on
enzyme present in the photosynthesis in this topic.
livingworld?Suggest Another group of autotrophic organisms are unable to use light energy. Instead, a specific
why. chemicalre-acrioniscatalysedinordertouansferthefreeenergyrequiredtosynthesise
organic compounds from carbon dioxide and water. Consequently, these organisms are known
as c hcmosynthc tic. Two important examples of chemosynthetic autO!rophs that we meet
elsewhereinthisbookare:
• the nltrlfying bac teria , that occur in the soil. lbese microorganisms o xidise ammonium
ions to niuites or niuites to niuates. Both of these reactions provide the bacteria with energy
needed to synthesis carbohydrates
• the iron bacte ria, which are sometimes used in industry in the e xtracrion of metal from
low-grade ores .
272
13.2 Investigation of limiting factors I
4? 13.2 Investigation of limiting factors
Environmental factor.; By the end of this section you should be able to:
influence the rate
of photosynthesis. a) explainth e te rmlimitingfactorinre lationtophotosyntho,sio
Investigating these shows b) explainth eeffectsofchangeoin light intensity, carbondioxideconce ntrationandte mpe rature on
how they can be managed the rate ofphotooyntho,sis
in protected environments c) explainhowanunde m a ndingoflimitingfactoni,usedtoincrease crop yie ldsinprotected
used in crop production e nvironments,suchasgl asshouse,
d) curyoutaninve stigationtodetermine the e ffectoflightinten,ityorlightwav.,l e ngthonthe
rate ofphotosyntho,si,uoingare doxindkator(e .g.DCPIP)and a suspensionofchloropluts(th e
Hill re action)
e) carryoutinvo,stigationsonthe effectsoflightintensity, a rbondioxide andtempe rature onthe
rate ofphotosyntho,sisusingwhol e plants, e .g.aquaticplantosuchME/odea andCabomba
olga,
collecting
~ tlary
:'u~!~te
Fig...-e13.14Measu,;ngthe,ateofphotosynthesiswithamiaob u,ette
273
IEJ Photosynthesis
,:;--- respi<alioo
o, I
( .)'°'
photosyothesi,
Then, as the light intensity increases funher, the rate of plxxosynthesis also increases; the leaf
becomes a net exponer of oxygen
Wecanseethegeneraleffeaofincreasinglightintensityontherateofphotosynthesisin
Figure 13.15. At low light intensities the rate of photosynthesis increase-sin proportion to the
increasing light. Here the low light intensity (i.e. the Jack of sufficient light) is limiting the rate of
photosynthesis. Light is a factor limiting the rate of photosynthesis under this condition.
However. as light intensity is raised funher. a point is reached where increasing light intensity
produces no funher effect on the rate of photosynthesis. Now some factor other than light is
limiting photosynthesis.
What may now be limiting the rate?
274
13.2 Investigation of limiting factors I
The limiting factor at that point is disclosed when the investigation is repeated at either higher light
(Figure IJ.16) or carbon dioxide concentrations, but not at the same time.
Look al tbisgrapb i>1 Figo,re 13.16now,mdfollow the annotations
Hi
i~~-; •
4 Yesl BPCaOsew!>entheexperiment isrepea!edata
~highe<CO,cOO<Pntratioo,therateolphotosynthe-sis
risesumilahigherlightimensityisreiKf>ed .
{""
iH;~ ,,..--
K!Wco,coocentratioo
2 Photosynthesisrateincrea"-"with
incrusing light imensity-Oight is limiting)
1 Wilhoutligllthere
isnophctosyothe,i,
lig ht intensity
Figure13.16Theeffectollightinlemityontherateofpootosynthesis
carbon dio~ ide concentra tion as a limiting factor The results of these earlier. physiological investigations fit well with our
h,cj, light intensi!y current understanding. Photosynthesis is a biochemical process involving a
,/
,-co,i,~not rate-lim~ing series ofintercotmectedrC3C1ions (Figure 13.19). All these reactions contribute
to the overall rate of photosynthesis. However, at any Ofle time, the rate
of phomsynthesis will be limited by the slowest of these reacrions - the
/ {/ lowlight intensity overall rate will be limited by the facror that is in shortest supply. This facror,
whichever Ofle it is, is known as the limillng facto r .
whatisnowlimitingther.Jte1 dearly,alimitedsupplycleitherlight(Figure IJ.16)orcarbondioxide
(Figure 1}. 17) could detennine the rate of photosynthesis. since both of these
are essential.
Both llglu inte ns ity and carbo n d iox ide con ccm rm io n can be limiting
factors in photosynthesis.
coocemra1iooofco,1arbitra,yunils
Figure13.17Theeffectofcarbondiox.Jeconc:entration
The effect of temperature on the rate of photosynthesis
on photosynthesis The effect of temperature on the rate of photosynthesis, under COfltrolled
laboratory conditions, is especially informative. This is because, one effect of
a t highlightintensitytemperntu,erise increasing temperature. say in the range from IO"C to 30°C. depends UJXXl
iocreasesrateofphotos;mhesis
lig ht inte n s ity. Under low light intensities, a rise in temperature has little
' effect. Under higher intensities, the rise in temperature increases the rate of
phomsynthesis significantly. This relationship is shown in Figure 13. 18.
Howcanlbiseffectbeexplait1ed?
/::c
A~:;:!,:
Under low light intensity, light is obviously the limiting facror. Now a light-
dependent reacrion, like all photochemical events. is temperature indifferent. In
.
the same way. a light-independent reacrion Oike all biochemical steps catalysed
by enzymes) is highly temperature sensitive.
lne graph in Figure 13.18 was interpreted as showing that phomsynthesis
------------------· ismadeupoftwosequentialstages.alight-dependentstageandalight-
independent stage. Later, this idea was confirmed. once phomsynthesis was
investigated by entirely different, biochemical techniques. including the use cl
chloroplastsisolatedfrornaleaf,for=ample.
IO 20
tempernture.l"C
Figure13.18Theeffectoftemperatuneonthe,ateofphotosynthesis
275
IEJ Photosynthesis
12 a Suggestwhywe
should expect
that high external
temperatures
cause the rate of
photosynthesis to
falloff very rapidly?
b What may
effectively prevent
over-heating of
greenleaves7
Figure13.19Photosynthl'Sis:asummary
The highly significant benefits d carbon dioxide supplementation may be offset to some =tent by
the =pense incurred in altering the growth environment in this way, and by the raised costs due to
t: t
the need lO monitor and o:introl the environment. For a:ample, infra-red gas analysers are needed
to monitor this environmental factor, linked to a computer to maintain pre-set. minimum and
maximum le vels of aunospheric carbon dioxide. The crop, concerned must be able to command a
premium price in nearby markets for this approach to be sustainable.
.
£
~
100
80
'~ 50
rnocentrationlppm
Figure 13.20Amb>eotCO,level,~ndnetphotosynthetic:productivity
276
13.2 Investigation of limiting factors I
Practical : Investigating the rate
of photosynthesis using an isolated
chloroplast suspension - the Hill reaction
Introducing the Hill reaction
O,loroplas1s can be isolated from green plant le-aves and suspended in buffer solution of the same
concemration as the cytosol (an isotonic buffer), see Figure 13.21. If the suspended chloroplasts are
undamaged, they function much as they do in the intact leaf. lbese isolated chloroplasts can be
used to investigate the splitting of water, for =ample. Howe ver, this requires the natural electron-
accepwr enzymes and electron-carrier molecules to be present, or the substitution of an alternative.
Jn the research laboratory, a sensitive piece of aPP"rarus called an oxygen electrode is used to
detecttheoxygengivenoffbyisolatedchloroplas1s.
Jn your laboratory, a h ydrogen -acceptor dye that changes colour when it is reduced is a
practical alternative. The dye, DCPJP (dkhlorophenolindophenol) is an =ample. DCPJP does no
harm when added to chloroplasts in a suirnble buffer solution, but changes colour from blue to a
colourless state when it is reduced. The splitting of water by light energy (photolysis) is the source
of hydrogen that turns DCPJP colourless. The photolysis of water and the reduction of the dye are
representedbytheequmion:
2DCPIP + 2H 20 ---> 2DCPIPH, +0 2
This demonstration is referred to a., the Hill reactio n after the research scientist who first
demonstrated this "reducing power" of Uluminated chloroplasts. It is a demonstration of the
light-dependentstageoutsideoftheleaf.
::·~~)
i=
,,,od,oa
medium(cold)
homogeniser
moto,base
;,i;;,tuge
Jfij_ ,o,"""'"'
~ p ellet
Jfij_ ,o,""'""'
~ p ellet
U--rnmb,oed
chloroplast
Figure13.21!so!atingintactchlorop!astsbydifferentialcentrifugabon
277
IEJ Photosynthesis
auffer DCJII'
(0.1%) (D.1 % sol)
Placedinthecolorimete,aod thescaleadjustedto
,ead0.7ootheabsorbancescale
Thenthetuberemoved.exposedtobright
light, andretur....dtothecolor;meterat
15secoodintervalsuntiloofurthe,<hangeinthe
absorbanceoccu"
Thistimeisrec:orded
Contents kept incomplete darkness{W<apped ;n
alumjnium foil) for the same period of time that
Tube1takestoshownolurthercolou,change
Thentfiefoilisiemovedaod theabsorbanceread
inthernlorimete,. andalso rec:orded
Thesetubesa,et,eatedinthesamewayasTube1.
maintainingtfiec)'Cleofexposu,eandrec:ord ingof
absorbanceforthesameperiodoftimeittakesfor
f - - + - - + - - + - - + - - - - - - - - < Tube 1 toshownofurthercolou,ch.lnge
Discuss the results}" " anticipate ,dtb µ,,ers and J<>Ur teacher, a'1d then ansiu,r Que.stio" 13
13 Given that a chloroplast suspension will decolourise a blue OCPIP mlution in the light,
explain:
a whythesteµstothechloroplastextractionwerecarriedoutinanice-cold waterbath,
where possible
b why,fOftheexperimentitself{tubesinthelightandthedark},thereac:tionmixtureswere
allowed to warm to room temperature
c whyanisotonicbuffersolutionisusedtore-suspendthechloroplasts,ratherthandistilled
d whatrnlourchangeyouanticipateinTubes1and2,ifany
e whatis!.Uesthecontroltubes(3and4}willrewlve
278
13.3Adaptations for photosynthesis I
13.3 Adaptations for photosynthesis
All the stages of By the end of t his section you should be able to:
photosynthe!.isoccur in the
chloroplast.Some tropical a) describe t herelationshipbetwffn•tructurea ndfunctioninthechloropla<tusingdiagramsand
crop1haveC,metaboli1m electron micrograph.
and adaptations to b) explain how the anatomy and physiology of the leaves of C, plants, such as maize or sorghum, are
maximise carbon dioxide adaptedforhighratesofcarbonfixationathightemperatur.,.inl<'rmsof:
fo@tion • thespatials10parationofinitialcarbonfixationfromthelightdepe ndentstage{biochemical
detailsoftheC,pat hwayarerequiredinout lineonly)
• the high optimum temperatures of the enzym"' involved
There is 3 double n:,cmbr:me around the chloroplast. The inner membrane inrucks 3t various
points, forming a system of branching membranes, called 1hylako id n:,cmbranes. These thylakoid
membranes are org:mised into many flat, compact, disc-shaped piles, called grnn., (singular.
gr:,.num). Between the grana are loosely arranged tubular membranes suspended in a watery
llllltrix, called the s trom3. Also present are smrch grains, lipid droplets and ribosomes. O,Jorophyll
pigmentsoccurboundtothemembrane.softhegram(Figure 13.22).
TEMoftha granumshowing
•
thyla koidm ambranas inwhich
chlorophylpigmentsa ra ha ld
(x38000)
olthegraoa
chlom!Jhyll pigmel\tsa,econtaioed
in!hegrana.s.,ndwichedbe1W""'11ipids
oodproteinsol!he thylakOOmembranes
Figure13.22Thestructureofthec:hlorop!ast
279
IEJ Photosynthesis
16 Oeducethesignificant
"""
difference between
the starting materials
Anatomy and physiology of C4 plants
andtheend-product5 The fixation of carbon dioxide in the Hgln-independeru step of photosynthesis was Ulustrnted
of photosynthesis in Figure IJ.11. page 270. We saw that carbon dioxide and the five-caJbon acceptor molecule,
ribulose bisphosphate. reacted together in the presence d an enzyme known as rubhrco, in
thestfOfllaofthechloropbst. Thefirstproducrisathree-carbonsugar, andbecauseofthis , that
pathway d rnrbon fixation is known as the Ci pathway. You ran see an Ulustration of how
rubis= functions in Figure 13.23 - and also the name of the three-carbon sugar product, should
you have forgotten it.
Photosynthesis evolved when the Eanh"s atmosphere had an e xtremely low concentration of
oxygen but a high concentration of carbon dioxide. This aunosphere was quite different from
our atmosphere today. Since those early days. the concentration of oxygen in the aunosphere has
increased (bemuse of photosynthesis!). You will be aware that today's aunosphere contains about
21percentoxygen.
Despitethesechanges,rubiscomostlyfunctionsnormally, particularlyingreenplantsof
temperate regions. However, in conditions =perienced by tropical and sub-tropirnl plarus,
thisisnotalwaysthernse. Plantsoftropicalandsub-tropirnldimatesexperiencemuchhigher
sunlightlevelsandhighertemperatures. lnsuchanenvironment,lightisnotalimitingfactorof
photosynthesis, but carbon dioxide often does become a limiting factor. in full sunliglu.
Jn these condi1ioru it has been discovered that oxygen from the air becomes a compctith'c
Inhibitor affecting the active site of rubis=. The unfortunate result is the breakdown of the
five-carbon acceptor molecule. ribulose bisphosphate, to a two-carbon compound. of Hnle value
to the plant (and one molecule of the normal three-carbon sugar product). This phenomenon
is known as phoiorcsp!rmlon . lt reduces the yield from photosynthesis. You can see this
competitive inhibition event of rubisco and the products formed in Figure 1}.2J.
280
13.3Adaptations for photosynthesis I
'\ /~-C:~;:,p1,o..
~ O-O~yg••
::::::---.._ pho,phoglycolo1e-isofli1tleuse10
~-......__,_c-ereoction,inchloroploshosit wasles
""-... + both ATP ond rad...cir.g powe< to rKyda
withribulosebi,pho,pho!aond02 otlhe C~-';,.
::;·~::i~t~~~~)':.f:'
molacule,olglyc"'ole3.pho,phola
~1)1,osphote
Figure13.23Rubisc:oinphotosynthesisar,dphotorespirahcm
The plants with this mechanism are called C. pla n ts, to separate them from all other green
plants Ulat fix carbon dioxide only by the Calvin cycle and produce a three-carbon sugar (triose
phosphate)asthefirstmainproducts - theCiplants.
C. plants have a different arrangement of the photosynthetic cells in their leaves. Here, the main
photosynthetic cells (called bundle s h c aih c ells) are surrounded by spcdal mcsophy ll cell"<
where carbon dioxide is fixed into a four-carbon organic acid (called malare).
So, in C.. plants in the light, the four-carbon organic acid formed in the mesophyll cells is moved
into the bundle sheath cells as soon as it is formed. In the bundle sheath cells the organic acid
breaks down, producing carbon dioxide and so ensuring its concentration is high (Figure 13.24). At
this concentration. carbon dioxide out-competes oxygen at the active sire of rubisco.
All the enzymes involved in the C.. pathway have high optimum temperatures and so funcrion
efficiently under ambient temperatures in the tropics, in daylight. As a result of this mechanism, C..
plants operate more efficiently at the high temperatures and high light intensities found in tropical
17 lnC 4 plant5,carbon regions of the globe. Many tropical grasses and plants of tropical and sub-tropical origins, such as
dioxideisf1xedinthe thecropplantssugarcane,maizeandsorghumareC..plants.
bundlesheathcell1a1 Sorsbum is another member of the grass family. We have already examined this plant as a
wellasthemesophyll xerophyte (page 143). It is a plant that grows well in hot, semi-arid tropical environments d high
cells.How does light intensities , where annual rainfall is typically only 400--000mm - conditions much too dry for
carbon dioxide fixation maize. In fact, Sorghum has become the fifth most important world cereal, after wheat, rice, maize
differ in these two
and barley. Like maize, it is a C.. plant. Examine carefully the features of the Sorsbum leaf that are
{adjacent)locat ions7
thesiteofC..carbondioxidefixation(FigurelJ.25)
281
IEJ Photosynthesis
-
re.ictionsofphotosynth~inpliotsof
temperateclimitesiscalledtheC 1 pathw•y
be<:.iluseth.lirstproduct11a3-arbon
Manyt,opicalplints(indudongSorghum)
R;Oproducei4-carboncompound(Mi
Of9i1111CKid.rNlite). Theyarekncwn6
"""""
ThestepSofC,pholosynthesisare
SU1T11Nnsedbelow,buttheeffectofC,
photosynthesrsistoinaNsethemiountof
COzt.1keninfromtMairwhiletMstomata
;i.reopff'I.Theproduct(malate)thenbreak.s
down;indrete-theC02 1odiMlionin thl'
~~:~=i~rt:~r:=.c~.. ::w
Figure1l.24 Thes1te.ondstepsofC,phoo::,syntha!i
181
13.3 Adaplations for photosynthe,is •
' indiOOH1$
-omorphic
feotvrH
proproots-
11r,ngrti.,,.d
wi1hoddirional
lib,..
•"e,aprion,lly""*'siffadvwuilousrOOllenwge
fromsllfflnodmjustbelowsoilie....
por1olld1«1ion {11S0J
•'lhey10p soil10ad,prholaboul l.8m,oodspr..:>d
lalefOlly1.Sm ...... main
.... scular upperepiM<mil
bundle wifhfflidwut:'(CUlidt
'ITlClffloirttoppedon
•• ul"Offlideoli.of-
- ~ slowswoltrYopour
lossbydiffutlonfrom
leafviostornolO
283
m Photosynthesis
• Photosynthesis is an energy trande r PfOCl'SS in which • A light-independent st age occurs in the stroma and
gr~ plants manufacture <arbohydrates from <arbon inwlYes the fixing of carbon dioxide to form trioses, lM1g the
diox ide a nd water, using energy from s unl ight . Oxygen productsofthe~ght-dependerltstage. Triosesareusedina
isthewasteproduct Chloroplasts,theo,ganellesinwhich varietydbiochemical reactions\\ithinthechloroplasttoform
photosynthesis OCC\KS. are chiefly found in the mesophyll cells rnono5aecharides(g!ucoseandfructose)thatareconverted
of the leaves, particularly the palisade mesophyll cells. Plants tosucrosefo,translocationo,tostarchfo,storage.Other
use the products of photosynthesis to manufacture the other reactions use minerals from the soil to form amno acids that
carbohydrates,lipids,proteinsand allothermetabofites, aretran51ocatedandusedfo,proteinsynthesis.
enzymesandpigmentsrequired. Fo,thistheyrequireenergy • lngreenplantsthevo!umeofo)()'gen--ennchedgasgiYen
transferredbyrespirationandselectedmineralions,absorbed off in the light is a measure of the r•te of photosynthesis
fromthesoil. Consequently.plantnutritionisdescribedas TheYOlumeofgasproducedbyaquaticplant:sunder
autotrophlc (self-feedin9). drfferentconditionscanbemeasuredusingamicroburette
• The lea f is the'faaory'fo,photosynthesis,withpali!.adecells (photosynthometer).
inpo5itionstoabsorbmaximumlight.Thecellsaresupplied • As photosynthesis consim of a number of interconnected
1, ithwaterfromthexylem vesse ls.andwithcarbondioxide reactions,theslowestwilldeterminetheoverallrate. The
fromtheairbydiffusionthroughopens tomata . The products factorlimitingitsrateisthendescribedasthe llmltlngfa ctor
ofphotosynthesisareexportedfromtheleafinthesieve Alimitedsupp!yofeithercarbondioxideorlight\\illlim it
tubes in the phloem. the overall rate of photosynthesis, sincebothareessential for
• Chloroplastsareenclosedbya doubl e membrane. The inner photosynthesis.
membran e intuch to form the membrane systems of the • Theautotrophicnutritionofthe greenplantalsornstains
chloroplast(thylakold membranes). These are arranged in other living things, sinceotherorganismsfeedonplantsor
compact, disc•shaped~lescalledg ran a,aroundwhichare plantproducts,directlyorindirectly.
!005elyarrangedthy!akoidmembranesofthe stroma.
• l'tants, induding crop plants show adaptations to
• Chlorophyll consists of a mixture of pigments, two of v.4-iich different environments. These include C, <a rbon dioxide
are chlorophylls. They can be separated by <hromatography. metaboli s m as part of photosynthesis in many successful
Measurements of the absorption spe<trum d chlorophyl and tropical and sutrtropical plants. This mechanism isa
thea<tionspectn.mofphotosynthesisshc:Mothatitisthe blue re5p00setotheharmfuleffectsofhighoq9en
and ,fit components of white ight that are selectively ab5omed conc:entratiom and loN carbon dioxide concentrations
andaremosteffectiYeinbringingaboutphotosynthesis on the carbon dioxide-fixing enzyme, rubisco. at high
• Photosynthesis can be divided into two. linked stages ambient temperatures. The structural adaptations sh<mn by
• A li ght -dependent stage occurs in the grana and results mesophyll tissue in the leaves of sorghum and maize provide
in the formation of reduced NAOP and ATP. It imdves the the venue for C. carbon dioxide fixation .
photolysisofwaterandthereleaseofoxygen.
284
Examination style questions I
Examination style questions
1 Aninvestigationintotheeffectoftemperatureontherateof 4 a) Theenzymerubiscoisthemostabundant, naturally
thelight-dependent1tageofphotosynthesiswascarriedout occurringprotein.Wheredoesrubiscooccur,andwhat
using isolated chloroplasts reactiondoesitcatalyse7Suggestwhyitsabundanceisto
Samplesofchloroplastssuspendedinbufferweremixedwith be expected [4]
arnlouredelectronacceptorandexposedtolight.Thernlour b)Byreferencetorubusco,explaintheprincipleof
changesfrombluetocolourlessaselectromaretakenupby competitiveinhibitionofanenzyme. [4]
the electron acceptor. c) The reactions of photorespirationreducetheyieldof
a) i)Sketchagraphtopredicttheresultsoftheinvestigation photosynthesis.WhyisthisMi7 [4]
12) d) ByreferencetoanamedC,plant,explain,withuseof
ii) ldentifytwokeyvariablesthatmustbecontrolled a diagram, thearrangementofcellsinarepresentative
inthisinvestigation.Foreachexplainhowitmight partofaleafthatisthesiteofC, metabolism. Annotate
be controlled [4) yourdiagramtoidentifythelocatiomofthestepstoC 4
metabolism. [8]
iii)Outlineaproceduretofindtherateofreactionforthis
(Total : 20]
investigation [2)
b) In a further investigation, small quantities of AOP 5 a) Describethestructureofphotosystemsandexplainhow
andinorganicphosphatewereaddedtotheisolated a photo1y1tem functions in cyclic photophosphorylation
c:hloroplastsbeforetesting (9)
Suggestanhypothesisbengtestedbythisfurther b) ExplainbrieflyholvreducedNAOPisformedinthelight-
investigation ltl dependentstageofphotosynthesisandisusedinthe
(Total : 9] light-independent stage [6)
(Cambridge International AS and A Level Biology 9700, Pilper (Total : 15]
51 QI November 2009) (Camtxidge lntemational AS and A Level Biology 9700, Paper
04 QIO November 2007)
2 a) Bymeansofalarge,fullylabelleddiagram,describe 6 a) Show how both the absorption spectrum and the action
thestructureofachloroplast,asdisclosedbyelectron spectrumofaparticularspeciesoffreshwateralgaemay
microscopy. [6] be measured simultaneously.
b)Annotateyourdiagramtoidentifythefunctionorroleof b)Drawandfullylabeltheabsorptionandactionspectra
thestructuresyouhavelabelled [4] youwouldexpecttobeproduced,usingasinglegraph
c) lninvestigationsofthephysiologyandbiochemistryof c) Describeinoutlinethestepsbywhic:hthechlorophyll
photosynthesis,rnspensionsofsingle-celledalgae{simple, presentinasampleofgreenleavescanbeobtainedand
photosyntheticorganismsfoundgrowing infreshwater the component pigments 1eparated by means of paper
environments}areoften1ubstitutedforwhole,green chromatography.
terrestrialplant1.0utlinetheadvantagesofthi1choiceto
the experimenter. [6)
(Total: 16]
285
A Level
14 Homeostasis
Ce ll s functionmos t e ffici e ntly iftheyue ke ptinnear con stant o f cell s.Priorknow ledge forthis topi c includ esa n
co nditio ns. Cell s in multice llula r a nima ls are surround e d by ti ss ue understanding th at wast e prod ucts a re exc reted from the body -
fluid. Th e composition , pH a nd te mpe ra ture of ti ss ue fluid ue a role th a ti s fulfille d by the kidn eys - a nd a n o utlin eofthe
ke pt con st a nt byexc ha ngesw iththe bloodas disc ussedinth e structurea ndfun ctio n o fthe nervousand e nd oc rinesyst e ms.ln
t o pi c o n Tra nsport in ma mma ls. In ma mma ls, core t e mpe ra tu re, pla nts. g ua rd ce lls respo nd t o fluctua tio ns in enviro n me ntal
bl ood g lucoseco ncentra tio n a nd bloodwate r pot e nti a l a re conditi o nsando pe n a ndcl osestoma t aasap pro pri at e for
ma in ta ined w ithin na rrow lim its to e nsure the e fficie nt o pe ra tio n ph ot osynth esis a nd co nserving water.
Homeosta1is in mammals By the end of this section you should be able to:
require1rnmplexsystems
to maintain internal a) discuM the importance of home ostasis in mammal, a nd expla in the principl es of homeortasis in
conditions near constant te rmsofint<'rna l a nd exte rnalstimuli,rec<'ptors,ce ntra lcontrol, co<>rdinationsy,tems, e ffe ctors
(muscl<'S andgland,)
b) de fine the t<'rm ne gat ive f...,dback and explain how it is involved in hom<'ostatic mechanism,
c) outl ine the roles of the ne rvous •Y"te m and en docrine syste m in co~rdinating homeostatic
mechanism,,indudingthe rmoregulation,osmo,e gulationandth e controlofbloodglucose
286
14.1 H~sinmammal, I
Maintaining a constant internal environment -
homeostasis
Living things face changing and sometimes hostile environments. Some =remal conditions change
slowly, others dramatically. For example, temperature changes quickly on land exposed to direct
sunlight, buc the temperature of wa!er exposed to sunlight changes very slowly.
How d o organisms ros/xmd lo e,wironmental changes?
Some animals are able to maintain their internal envirorunent, keeping it more or less constant,
allowing them to continue normal activities, at least over quire a wide range of extern:il conditions .
These are the ' r egulatono' . For example, mammals and birds maintain a high and almost constant
body temperature. Their bodies are kepi at or about the optimum temperature for the majority of
the enzymes that drive their metabolism. Their muscles contract efficiently and the nen,ous system
co--ordin31es responses precisely, even when external conditions are unfavourable.
Regubtorsaredtenabletoavoiddangerandtheyrn:,yalsobenefitfromthevulnerabilityof
prey organism., which happen to be ' n o n .n,gula tors' . So regulators rn:,y have greater freedom
in choosing where to live. They can exploit more habitats with differing conditions than 'non-
regulators' can, too. But whether an organism is a 'regulator' or nOI:, some of control of the internal
env· ment 'sessent' J.
llo m costas is is the name we give to the ability to maintain a constant internal environment.
Homeostasis means 'staying the same'. Mammals are excellent examples of animals that k=p
their internal conditions rern:,rkably constant. Their internal environment is the blood circulating
in the body and the fluid circulating among cells (tis.sue fluid) that form, from the blood plasma,
delivering nutrients and removing waste products whilst bathing the cells. Mammals succes.sfully
regulate and maintain the pH, the concentrations of oxygen, carbon dioxide and glucose, the
temperature and the water content of their blood. All these are rn:,intained at constant levels or
within very narrow limits .
How is bomeoslasis brought about and maintained'
pHofblood
Mamm als are a comparitively recent group in terms of
the ir evolution aryhistory, yetthey havesuccessfully
settledin s ignificantnumbe rsinvirtually ev e rytype
ofh abitatonEarth.This successisdirectlylinkedto
the ir abilityto controltheirintern al e nvironmentby
Figure 14.1 Homeostasis in mammal, home taS s.
287
e:J Homeostasis
componentsofanegativefeedbackcontrolsystem
input- ~ receptor ~ co.ordinator ~ effector ~ outpul-
<hangero m&ISlXP<"-"'el levejo/operatiooisse!he<e,aJ>d b<ingsaboutasecood rondilion<P<torl'd
!he.lJ"!>'m al the variable l1formatioofmmrec:eptorll'O'MedaJ1d change to system toserva/ue
romparedlMrhservalie,aJ>d mmmands Onoppositedi"eaion
+ roeffemx despatc:hedlmmhere rothel1pt1t)
the laboratorywaterbathunit,aneumple
ofaself-regulatingsyitem
c0-0rdir1,11orw~h
onfolfsv.itch, pattemofchangetowaterbathtemperature
:'!"'!-point (waterba!h mrnmlsetat25"C)
i 25 -
±i
I w evenwithaho~lyeffi<iem
I 15
rer:eptorde"°'itis
impossibietoa\Oidsome
'o""rshoot'inregulation
Figure14.2Negatrvefeedback,themer:hanism
288
14.1 H~sinmammal, I
Homeostasis in action - control of body
temperature
Figure14.3Kowheatis
traf"!'iferredbetweenorganism
and environment
avaporation-the
c:hangeolllquidto
vapour - accomp.a,niedby
~::.t:.~=:.~
roolingeffect
CondocUoo.ritdiatioo
~';;.~::::toor
ofan"'gani,m
conductionofhutlromthehonerto
!hernlde<oftwo,u,!ac:esinrnmact
The regulation of body temperature, known as thc rn10rcgula1lon , involves controlling the
movement of heat across the body surface. l>fammals maintain a high and relatively constant lxxly
temperature, using the heat energy generated by metabolism within their lxxlies (or by generating
additionalheatinthemusdeswhencold)andcarefullycontrolHngthelossofheatthroughthe
skin. An animal with this form of thermoregubtion is called an c ndoihcrm . meaning 'inside heat'.
Humans hold their inner body temperarure (core temperarure) just below 37°C. In fact, human
inner body temperarure only varies between about 35.5 and 37.0°C within a 24 hour period, when
weareingoodheahh.
• by the sweat glands, which produce swem when the body needs to l = heat, but do not when
the body needs to rcrnin heat.
289
e:J Homeostasis
roleofthesweatglandsinreg ulatingheatloss
through the skin
sw,,atsecretedoo ;nb'>'hu,,.;c1;1y,.,..d
ms.kio,urfacp \;:"',,;ngai",wapo ration
\occu,sandcau5l".Scooliog
T
skiocapillary
~H-- -:;:::=·
roleofcapillariesinregulatingheatlossthroughtheskin
roleofthehairinregulatingheatlossthroughtheskin
Figure14.4Thes.kinandtemperature,egulatioo
290
14. 1 Homeosla!is in mammals I
elsewhere that decrease heat loss (such as vasoconstriaion d skin capUlaries) and increase heat
production (such as shivering, and enhanced 'brown fat' respiration). When body temperature is
higherthannonnaltheheatlosscentreinhibitstheheatgaincencreaaivity, and sends impulses
tothe skin, hair erector muscles, sweat glands and elsewhere that increase heat lo.ss (such as
vasodilationdskincapillaries).anddecreaseheatproducrion.
~ ---+-=::==1~
I
, , po,ili<,n'™"'911'(1
291
e:J Homeostasis
• llypoglycaem.ia. in which our blood glucose falls below 2.0rnWL. If this is not quickly reversed,
we may faint. If the body, and panicularly the brain, continue to be deprived of adequate glucose
levels. convulsions and coma follow.
• llypcrglycem ia, in which an abnormally high concentration of blood glucose occurs. Since a
2 Explainwhatlivercells high concentration of any soluble metabolite lowers the water potential of the blood plasma,
receive from blood from water is drawn immediately from the cells and tissue fluid by osmosis, back into the blood.
thehepaticarterythat As the volume of blood increases. water is excreted by the kidney to maintain the correcr
is not present in blood concentration of blood. As a result, the body rends to become dehydrated and the circulatory
from the hepatic portal system is deprived of fluid . Ultimately, blood pressure cannot be maintained.
For these reasons, it is critically important that the blood glucose is held within set limits
bloodglliCose serumimulin
mmolllitre pmolilitre Regulation of blood
- - - - - - - - - - - - - - - - - - 400 glucose
- bloodglurnselevels Regulation of blood glucose is
- blood insul in levels
) theresultoftheactioruoftwo
= 'ilarch-richfood hormones, released from groups
ofspecialcells,foundinthe
=== rucrose-richfood
pancreas.
After the digestion of
carbohrdratesinthegut,glucose
is absorbed across the epithdium
cellsofthevUH(finger-like
exrensionsoftheinnersurfaceof
the small intestine) into the hepatic
portal vein(Figure8.7 , page 158).
The blood carrying the glucose
reachestheliverfirst.lftheglucose
level is too high, then glucose
scans to be withdrawn from the
50 blood and is stored as glycogen
However. notalloftheglucosecan
3.0 e-~~-~~-~~-~~-~~~-~ be removed immediately. Blood
7a.m. 9a.m. 11a.m. lp.m. 3p.m. Sp.m. 7p.m. 9p.m. 11p.m. la.m. circulating in the body immediately
afteramealhasaraisedlevelof
i breakfast i 1unch t dmoe, glucose(Figurel4.6)
292
14.1 H~sinmammal, I
At the pancreas, the presence of an =cess of blood glucose is detected in patches of cells known
as the Ls,lc1 s of l.angerh a n s. These islets are hormOfle-secreting glands (endocrine glands); they
have a rich capUlary network, but no ducts that would carry secretions away. lnscead , they are
transported all over the body by the blood. The islets of I.angerhans contain two types of cell.
a lp Im ( 11) cells, and beta (13) cells (Figure 14.7).
Figure14.7Anis!etofLmgeffiilnsinthepar,cn,as
Jn the presence of a rnlsed blood glucose l~-vcl, the bc1a cells are stimulated. 1bey secrete the
hormOfle ins ulin into the capillary network . Insulin stimulates the uptake of glucose by cells all
over the body, but especially by the liver and the skeletal muscle fibres. Anocher effea of insulin is
to trigger the conversion of glucose ro glycogen (glycogcncsis) and of glucose to fany acids and
fats in liver cells. Insulin also promotes the deposition of fat around the body.
3 Giventheroleofalpha As the blood glucose level revens ro normal this is deieaed in the islecs of Umgerhans. and the
cells and beta cells beta cells respond by stopping insulin secretion. Meanwhile the hormone is =creted by the kidney
(the production and tubules and the blood insulin level falls.
discharge of hormones), When the blood gluco,;c level falls below n ormal , the alpha cells are stimulated. These
identify the organelles secreie a hormone called glucagon. This hormone aeiivates the enzymes that conven glycogen and
that are involved
amino acids ro glucose (glucon cogcncsls). Glucagon also reduces the rate of respiration.
directly and list their
As the blood glucose level revens ro normal. glucagon production ceases, and this hormone in
specific roles
turn is removed from the blood in the kidney tubules
pancreas
cootainsspecialcells oftwo
type,(illtheisletsof
< ~ ~~~~harn) that are sensitive
ose ~levelol,ugarilltheblood
·~ :
-lli..,.!,.ii' :~·i~glucose
bmdoootex""'1~
in,ulincJ sesklwe,ir,g
ol bloodglucose~ve!
?glucoselevel
~~=
Fig..-e14.8Distriblllionandmetabolismofbkxxlglucose
293
e:J Homeostasis
Figu,..,14 .9GltKoseiegulaboo
bynegatrYeleecbad
294
14. 1 Homeosla!is in mammals I
The impact of adrenalin on liver cells as a three-stage process
in.Ktive kinaseeozyme-G,p
of cell-signalling
act""'kinaseenzyme The impact of adrenalin on a liver cell is a 1hrcc-stagc p rocess of cell signalling
inactivephosphorylasel'l\zymet (Figure 14.11). After the first stage, when a honnone moleo,le interacts at the cell
surface,subsequenteventsareactivatedbyasuccessionofrelaymolecules,initiatedby
the formation of a second messenger, cAMP, within the cytosol.
,;,,c.,:i"°"""'""ffi"""' The relay molecules are activated kinases and phosphorylases. These proteins
catalysespecifi.ctypesofreaction·
• Kinascs are enzymes that transfer a phosphate group from ATP to an accepior.
glucose1-ph05pflate
• PhO!!i phorylascs are enzymes that break down glucose-based polysaccharides (e.g
Figure14.11Thethreestagesofcell-,ignallir,g
glycogen)toglucose I-phosphate
that;,dm.,lintfiggers
The critical role of the second and third stages of signal transduction is the amplifica tio n of the
hormone signal. So, from one activated recepior molecule, 10000 (10') molecules of cAMP are
formed. As a result of the presence of cAMP in the cytosol, approximately to" moleo,les of active
phosphorylase enzyme will be formed, and these will then trigger the formation of perhaps to"
moleculesofglucosel -phosphate.
Table14.2Stagesofsignal Stage Conse quences
transductionleadir,gto
ampl iftcaboo hormoneinterac:bonat thecellsurface asingJehonnoneinitiatesevent,
format.in of cydic AMP. whITTl binds to l<inase prnteins about 104 molecules of cAM P become
active inthecyt050I
anl'l\zymecasc:i>deinvolvingac:tivabooolenzymesby apprnximately10 1 molec:ulesofglucose
phosphory!at.in to amplify Ifie signal 1-phosphate are formed and become
avaHabJetopassoutintotfiebloodstream
Type I diabetes (insuli n-dependent diabetes, early Type lldiabetes (d;abetesme llitu,. la te ooset diabetl'5
onsetdiabetesorjuvenilediabetes) oradult onsetdiabetl'5)
affect,;youngpeople,beklwtheageol20years the commoo form {90% of all cases of diabetes an,
of thistwe)
duetothede5tructkmofthe~cell1of the i1\et,;of
co mmon inpeopJeOYer40yearsespeciallyil
l.angerham by t!li' body's own immune system OYerweight. but thisloonofdiabetesisha,..;ngan
symptoms:
-constantthirst
- undiminished hunger
,,..,._ ..;:s,;,i =::::: .:.tt:~;:;::;,::::t:: ~~~~~nt:=~
countries.seem ir,glybecauseofpoo,diet
-excl'51iveurinatioo '".l!lP1~- . . , . * symptoms:
treatment: mikJsufferersusuallyha..., sufhcientbloodinsulin. but
insulinreceptorsoocells ha..., becomedefective
- iniectKlflofinsuli n intothebloodstn>amdaily
treatment:
- regular meil'>urementofbloodglucoselevel
la,gelyby dietalon e
295
e:J Homeostasis
Glucose is nOI: present in the urine of a healthy person because. although it appears in the
filtrareintheBowm3n'scapsuleofthekidney,itisselectivelyre3bsorbedintheproximal
convoluted tubules (page 301). However, people with dia.betes rnnnot control their blocxl sugar
levels effectively. Glucose may appe3r in the, urine when the blocxl sugar levels rise sieq,ly,
for example after a meal. Dia.becia; may need to inject insulin to reduce blocxl sugar levels. The
urine may be tested to find out if an injection is needed using dinistix'". Alternatively, a glucose
biosensormaybeused(seeFigure 14.13).
lbe dini.stix'~ Slrip conmins two enzymes, glucose m:idase and peroxidase, together with 3
colourless hydrogen donor compound called chromogen. When the strip is dipped into the urine
sample, if glucose is present it is oxidised to gluconic add 3nd hydrogen peroxide. lbe second
enzyme cataly""" the reduction of hrdrogen peroxide and the oxid3tion of chromogen. The product
is water and the oxidised dr,,, which is coloured. The more glucose present in the urine, the more
the principles coloured dre is formed. lbe colour of the test strip is then compared to
gltKoseoxidase theprintedscaleroindicatetheamouruofglucoseintheurine
glucose+oxygen - - -- gluconKacid+H1~ (figurel4.12).
An3hernativeapproachforthedi3beticistomeasuretheglucoselevel
pernxklase intheblocxlitselfusingagluc,-,blo,;en..-.or (Figure14.13).Abiosensor
DHi+H.,Oi 2Hi0+0
is a device which m3kes use of a biological molecule (or sometimes 3
lf
redtKed duomagen cell) to detect 3nd measure a chemical compound.
d1fomagen (coloured) The glucose biosensor has an immobilised enzyme. glucose o xld a.sc,
{colourless)
held between two membranes pasitioned at the tip of a platinum
electrode. In use, the outer membrane is momentarUr brought in contaet
with a tinr drop o f blood, squeezed from a pinprick puncture of the skin
<'• e•o«n at the tip of a finger. In contact with the immobilised enzyme, glucose in
test,1,;pd;pped ·- the blood plasma is irnmediatelr oxidised to gluconic add and hydrogen
rntoufinesample peroxide. The electrode measures the drop in oxygen used to produce
the hrcJrogen peroxide and an electrical signal is generated. lbe size of
thissignalispropon::ionaltotheconcentrationofglucoseinthepatient's
Figure14.12Mea5uringglucoseinurineusingaClinisfo::™ blood. A digital read-out gives the concentration of blood glucose.
~:;11-itj
lay<1r ' •
ratorbonam .
296
Figure14.13Q'°""""'forbk>odglucosetesting
e
14.1 Kidneys-structure mdfuiction I
14.1 continued ... Kidneys - structure
and function
- ~ excretedbytfie
kidneys
297
e:J Homeostasis
Associated with =cretion, and very much pan of it, is the process of osmoregulation.
Osmoregulation is the maimenance of a proper balance in the water a nd dissolved substances in
the blood. Excretion and osmoregulation together, are the work of our kidneys.
Figure14.15Thehumanurin.:uysystem
In section, each kidney consists of an outer con e x and irmer medulb , and these are made up of
a million or more ncphron..... A nephron is thin-walled tubule about 3cm long, pan in the o:irtex
and pan in the medulla. The shape of a nephron and its arrangement in the kidney are shown in
Figurel4.16
Blood vessels are closely associated with e ach of the distinctly-shaped regions of the nephrons.
For example. the first pan of the nephron is formed into a cup-shaped renal ca p s ule and
the capillary network here is known as the glo m crulu... Collectively, these are known as the
Jl.la lplghian body. They occur in the con=. The com ·olutcd tubules occur panly in the conex
and panly in the medulla, but n(){ice that the =tended loops of lien.le and collec ting duels
largelyoccurinthemedulla.
Each region of the nephron has a specific role to play in the work of the kidney. and the
capillary network se1Ving the nephron plays a key pan, too, as we shall now see.
298
14.1 Kidneys-structure mdfuiction I
n• phron w lthblood ca pllla~u
Photomicrograph o f the tortu of th• kidney In settl on. , howlng the rubules. r<1 nal capsules a nd capill ary networks
di stal
""'
~~~:::~.d
tubul es
Figun,14.16Thekidneyand,tsnephron,:rtructu,eandrok-;
299
e:J Homeostasis
The blood pressure is raised at this point by the input capillary (afferent arteriole) being
wider than the output capillary (effe rent arteriole) . This increase in hydrostatic (blood) pressure
exceeds the water pcxential of the plasma . This forces water and the soluble componems of plasma
that are able to pass out through the a:tremely fine sieve-like wall structure here, between the
podocytes, into the capsule. This 's kve · is made d two layers of cells (the endothelium d the
capillaries d the glomerulus and the epithelium of the capsule), between which is a basement
5 a Whatisthemurce membrane. You can see this arrangement in Figure 14.17.
of the force that Notice that the cells of the capsule wall are called podoq1es for they have feet-like =tensiOfls
drivesultrafiltration thatformanetworkwithtinyslitsbetweenthem.Similarly, theendotheliumofthecapillarieshas
intheglomerulus7 pores , too. This detail was discovered using the electron micro.scope, because these filtration gaps
b listthemain are very small indeed. So small. in fact, that not only are blood cells retained, but the majority of
components of the
blood proteins and polypepi:ides dissolved in the plasma also remain in the ciK"Ulating blood. It
blood that are likely
is the presence of the basement membrane that stops large prot.eins passing out. So. the fluid
tobefilteredinthe
nephron. that has been filtered through into the renal capsule is very similar to blood plasm.a but with the
significantdifferencethatproteinmoleculesarelargelyabsent
capilia<ies of
theglomerulus
-
(h~blood "
p<e<sull'here)
,_
wajlofll'<lill
epithelium) m,deu,of
::;""''
podocy1..
Fi g u re 14 .1 7Thesiteofultrafi!tration
300
14.1 Kidneys-structure mdfuiction I
Step 2: Selective reabsorption in the proximal convoluted tubule
The proximal convoluted rubule is the longest section of the nephron and it is here that a large
part of the filtrate is reabsorbed into the capillary network. lbe walls of the tubule are one cell
thick and their cells are packed with mitochondria. (\Ve would expect this, if active transport is pan
of the way re3bsorption is brought about.) lbe cell surface membranes of the cells of the tubule
wall in contacr with the filtrate all have a 'brush border' of microvillL lbese mkrovilli increase
6 a Cellsofthewalls enormously the surface area where reabsorp1ion occurs. The mechanisms of reabsorption are:
of the proximal • :,ctlve trans porl of s ugar., and amino acids across the cell surface membrane by the activity
convoluted tubule o fspecialcarrierproceinsinaprocessknownasco-transpott. lnthis.thecarrierproi:einusesthe
have a brush border diffusiOflofhydrogenions(prorons)downtheirelectrochemicalgradientinrothecell todrive
{Figure 14.19) theuptakeofmoleculesofsugarssuchasglucoseorsucrose.typicallyagainsttheirconcentration
Explain why this gradient. Other essential membolites are transported similarly.
feature is helpful • movement of mineral ions by a combination of actfrc transport. facill1a1cd diffusion and
totheproce-;101 somecx d.,gof·ons
reabsorption .
• diffu s ion of urea
b Whatdoesfacilitated
• movement of proteins by plnocytosls
diffusion involve?
• some movement of water by oisn10sis
Theelectmchemical9rad""1tinWions
betweentheextefio<andtheintefio<of
thecelldtivestheactivetransportof
metabolites{e.9.s.uga".amoniacids)
acms, the membrane a, the W flow
down their elernochemical gradient via
theco-transporte,pump
Figure14.18Co.\ransportact,Ve
trafl<portdtivenby a concentratioo
gradient
~~::;:~:...
backimobiood
bypino<ytosis
Figure14.19Reabsorptioninthep,oximalconvolutedtubule
301
e:J Homeostasis
::::::ri:IO
rubt,H,wall,
adjaceru3SCCOdinglimb.Asthe
filtrnteflows.theconcerurating
effeaismultipliedandsothe
fluklinandaroundthehairpin
water loss causes the benddtheloopsdHenleis
~-
Ahighcoocentratioool~tsislormedintl>emeli.Jl a. solutionolioosto
whi::ha llowswall'<!Obeabsorbedlrnmthenea<by becomernoa.>nlfated
collectingdoct,
Figure14.20ThelunctKlf\ingofthekq,olHen~
302
14.1 Kidneys -structure mdfuiction I
The role of the vasa recta is first ro deliver oxygen to and remove carbon dioxide from the
metabolically active =lls of the loop of Henle. NJ it does this. the blocxl in the vasa recta also
becomes saltier as it flows doum beside the ascending limb and Jess salty as it flows back up and out
7 Whati1theessential
featureofacounter- o f the medulla . In this way the cells of !he loop are serviced without remming the accumulated salts
currentflowsystem? from !he medulla. The vasa recta does absorb water that has pas.sed into the medulla at the collecting
duas. We discuss the working of these ducts in step 5. on the n=t page.
Osmon,gulation. the control of the water coment of the blocxl (and therefore of the whole
body) is a pan of homeostasis -- another example of regulation by negative feedback.
Howexaclljiistbisbrougbtabout?
The hypothalamus, pan of the floor of the forebrnin (Figure 14.21). controls many lxxly
funaions. lbe composition of the blocxl is cominuously monitored here, as it circulates through the
capillary networks of the hypoi:halamus. Data is also received at the hypothalamus from sensory
receptors located in cenain organs in the body. All these inputs enable the hypothalamus to
accuratelycomroltheaaivityofthepituitarygland.
The pituitary g land i, situated below the hypothalamus, but is connected to it (Figure 14.21).
The pituitary gland as a whole produ=s and releases hormones (it is pan of our endocrine system.
see below) - in fact it has been called the master hormone gland. In the process of o.smoregulmion,
it is the posterior part of the plmilary that stores and releases antidiun,tk hormone (ADIi) -
amongst others. (Other pans of the pituitary secrete hormones regulating a range of other body
activitie.sandfunaions .)
303
e:J Homeostasis
Aneumsecrei:orycel
ol!heh)POthalamus
"""""'"'"~•-o<cm
.:)·F-.'.~
,,
"'--;
,
acoonpotential
axoo
cel body
=======-00 000
ner,,:,e•nding
::::;:;..<apilla,ywall
~A; - · - · " ' "'
c"!)i lla,y
The cell surface membranes o f the cells that form the walls of the collecting ducts contain a
high proportion of channel pro{eins that are capable of forming an open pore running down their
centre. You can see the structure of a fluid mosaic membrane and its channel pro{eins in Figure 4.J
(page76)andFigure4.9(page82)
Wh~·n ADH ls p resent in 1.hc b lo od circulating past the kidney tubules. this hormone causes
the protein channels present in the collecting duct cell surface membranes to be open. As a
8 a List the components result, much water diffuses out into the medulla and very little diffuses from the medulla into the
of a negative collecting ducts.
feedback~tem Ca>l you e::,.p/ai>1 u·hy? You may "eed to go back lo sl,p 3 aboi,e.
andidentifythese
romponentsin The water entering the medulla is taken up and redistributed in the body by the blood circulation.
the process of Only small amounts of very concentrated urine are formed. Meanwhile, as ADH circulates in the
osmoregulation blood, the actions of the liver continually remove this hormone and inactivate it. This means that
b Howistheeffect thepresenccoffreshlyreleaseclADHha.s a regulatory effect.
ofADHonthe Wh~·n ADH is absent fro m the b lood circulating past the kidney tubules, the protein channels
collecting ducts
in the collecting duct cell surface membranes are dosed. The amount of water that is retained by
fed back to the
the medulla tissue is now minimal. The urine beccme copious and dilute.
co-ordinator7
9 Whyi1 toomuch
water rarely a problem
forplantcells,but
Urine analyses in medical diagnosis
potentially hazardous to Ananalysisofaurinesampleforthepresenceofglucose,pro{einorkeronelxxlie:sisan aidtothe
animalcells7 early diagnosis of serious but treatable conditions, as summarisal in Table 14.2.
Table 14.2ThevalueoluririeaMly;esinmed,c,Jldiagnosis
Tested for by
gltxose Clinistix',.orb,0Sen1or Presenc:eofglucosemaybeaniOOicatorof diabetes
protein dipsticktestforaliumin Presenc:eofpmteini1anindicatorof prote inura - apo,,;ible
indicatorofkidneydisea1ea,isingfromglomerulamembrane
damage.orseverehypertensioo
dipstick test - Ketostix"" Ketooes are a normal product of fatty a6d metaboli1m. but are
notoorma llypresentintheuririe . However.ifthebodyis1hort
o f gluco1e.thenfat1areuseda1anene ,gysoun:eandketones
may thena r,pea,intheufine.Typica llythisistheresultoltype
1diabetes.oradva nced/ untrea tedtype2 dia betes.but
eatingdi sord ersa rear,otherpossiiil ity
305
e:J Homeostasis
• drunkalmofwater • 1akeni11linlewater
th<>hypmhalamusdetectsthi,andstopsthe o sw,,atedexcessive!y
posteriorpituita,ryglar.dsecretin,g ADl-1 • eaten salty food
thehypothalamusdetemthisanddirl'Clsthe
poste,iorJ)ituitaryglar,dtosecreteADH
co rtex
m«lulla
wate,staysin
~rollediog
:~'',,'""'"=\
+ medultaabsomsless
thirst~<ITT - nofeelin,gofthim, water lromthe
/
=mcmw"• =,rn,,,,., """""""=• '
::.~~=sa~,}
(become,less""9alive) ',
larg e volume
of dilute
',, """" '°':""
( ', '.
blood rlasma~\tion- ~ -- ADHpreseminplasma ~ - - blood:lasma~tat·on-
' '
corrl'Clwaterpotemial('I'} is excrete d in kidney; rorrect""'mpoternial( ,p)
/
,' '
,," small volume
roo/irr/ewarer ,," of concentrate d
::.~=•hdsal!) ,, . f «I
306
14.2Horieostasisinplao15 I
14.2 Homeostasis in plants
c) describe the role of absdsic add in the d0<ure of stomata during times of wat~ stress (including
th eroleofcald umionsasasecondmessenger)
307
e:J Homeostasis
lnanexperimentaldemoostrabonttlallurg<lf Observation:onreleaseollhetu,gorJ)fe=re
P""...-eoltheguardceWscausestheopening inonegua,dcell,thedisbnc:liwshaJlf'Olthe
of the stomatal JK"", a mkmdi=!ion neecle cellwhenthepon,isopenwaslost.'Half'of
wasinsertedintoagua,dcel thepon,dlsappeiln'd
Befo<elheexperimentthe
stomataipo<ewaslullyopen, __
~;,to theturgornllhegua,d
Figure14.25Theopeningofstomata it'sturgo,pn,ssurethatdoesitl
apertu re
may vary
308
14.2Horieostasisinplao15 I
The accumubtion of these substances in the guard cell vacuole muses the ,vat er potemfal there
to beccme more neg:11ive. So, net uptake of waler from the surrounding ordinary epidermal cells
occurs - makingtheguardcellsextremelyrurgid(Figure 14.27).
dOllingisbroughtaboutbythereversalof thesesrepsinthedark. Alternativelyclosureoccurs
w hen triggered by the scress hormone absdsic add (ABA), produced in leaf cdls during w ilting. We
willdiscussthissitu31ionnext.
lnthe llght
lorilwo19rslfessdevelopsd ue 10wihing)
....,.,,.. w;l~ngofleof
producasflaccidaill,
ondlherefore
-. ou10moticS1oma1<1lclosura
call,underwotar
membrane around m u slsligh1wihingl
produceAIJA
V<Jcuolepumps
K~ ions in from
epidermal calls
- --4+"8-.f- K~ ionspumpedou1
in1heguordcalls, olvocuoletoolher
Sloredstorchis apidermolcells
coovon&d1oorgonic
acids, e .9.malo111, =',-Jl~~;;;;Jil;;;:;!
which are pumped
inlDV<ICUOle
molo1epumpedbackin1ocytoplasm,
cooverl8d10sugor,andstoredasstorch
Fig ure 14.27Howopen ir,gar.dclosir,gof,tomataoc:cu"
309
e:J Homeostasis
c) Howdostre11conditionswithinleavesduetowater
,~
from the body. [9) shortageaffect1tomatalopening,andwhatpartdo
[Total: 15] plantgrowthregulatorsplayinthisresponse? [7)
(Cambridge /ntemationa/ AS ;md A Level Biology 9700, [Total : 20)
Paper 04 Ql November 2005)
4 a) ldentifythedistinctiverolesoftheendocrineand
2 a)Afterthedigestion ofcarbohydratesinthegut,glucose nervom syo;tems in homeostasis, illustrating your answer
isabsorbedintothebloodstreamandtheconc:entration by reference to the mechanism of thermoregulation in
ofbloodglucoseinitiallyrises.Describehowa!.Urgein mammals [10)
blood glucose is normally regulated by the body and b)Outl inetheroleofa'secondmessenger'intheimpactof
whathappemtothebulkofthismetabolite a peptide hormone such as adrenalin on a liver cell. [10)
b) Draw a flow diagram showing the steps by which the [Total : 20]
concentrationofbloodgluco5ei1heldconstantbya
processofnegativefeedbi!Ck
[Total: 20]
310
A Level
The nervous system By the end of this topic you should be able to:
providesfastcommunK".iltion
between receptors and a) compare the nervous and e ndocrine system, as communication systems that co-ordinate responses
effectors tochangesinthe intemalandexte<nalenvironment
Transmission between b) describe the structure ofas=soryne urone andamotorneurone
neuronestakesplaceat c) outline the rolesof..,nsoryrece ptorcellsinde te ctingstimuliandstimulatingthe transmissionof
synapses n~·mpul-·nsensoryneurones
d) describethe functionsofse nsory,relayandmotornf'uronesinarf'flf'xa,c
<') describf' andexplainthe transmissionofanactionpote ntialinamyelinatednf'urone andit.
initiationfromare,<!ingpotf'ntial
f) explaintheimportan<f'ofthe myel insh..ath(sa lutatory<onduction)indetf'fffliningthf'speedof
nf'rveimpulSe5andthe,efractoryperiodindeterminingthe irfrf'qu<'ncy
g) des<ribethe structurf' ofacholine ,gksynap..,and explainhowitfunctiom,indudingthf' rolf' of
h) outline the roles of synapses in thf' ne rvous ~ e m in allowing transmi .. ion in one dirf'ction and
inallowingconn«tiombetwef'none ne uroneandmanyothers
311
m (onttol and CO·Ordioation
co-o«linatioo
'"""T"''
Offlpulsestransmitted
"'' ~ ' " ; ~ ~ ~ ~ = oftrn> I
j
effecroragans
,/ ''--..
musk-; ~
feedb.Kk
jnformarion
'"'Pllfl"-" - - - j
312
15.1 Control and co-ordination in mammals I
Surrounding the neurones there are differem types of supporting cdls called glia cells. lbese are
also an impormm pan of the nervous sySlem. In the brain and spinal cord there are several types of
glia cells. Jn the peripheral nervous system, the axons of many neurones are endooed by glia cells
called Schwann cell~. These wrap themselves around the axon with many byers of cell surface
membrane, forming a structure called a m yclin s heath (Figure 15.2). lbe sheath consists brgely
of lipid with some protein. Between each pair of Schwann cells is a tiny, uncovered ju oction in the
myelin sheath, called a node of R:m,·kr. The myelin sheath and its junctions help increase the
speed at which impulses are conducred - a point we will return to shortly.
r<tlay neurone
many fibres
di"ectioo
"'
impulse myel insheath
cell body
Figu~15.2Neuronesoftheriel\lOllssystem
313
m (onttol and CO·Ordioation
-=~"'- <=i;";::...""
.
p«'SSUfe
I
+"~~"'- efll'Clor
(e.9. mu«:le)
Figure15.3Thela)'OUtofareflexarc::thestructuralbasisofreflexactKl11
Howdoesarejlexarcu'Ork?
The action begins when a sense organ detects a stimulus. which is a form of energy such as
sound, light or mechanical pressure. This may become an impulse transmitted by a neurone that
servesthatsensecell.Oncegenerated,theimpulseistransmittedalongdendronsandaxonsof
asequenceofneuronesofthereflexarctoaneffectororgan. When itarrivesattheeffector,the
impulse causes a response - for example, it may cause a muscle to contract or a gland to secrete.
314
15.1 Control and co-ordination in mammals I
Sensory receptors and the conversion
of energy into impulses
Allcellsaresensitivetochangesintheirenvironment.butscn seccllsarespecialisedtodetect
stimuli and to respond by producing an impulse (an ae1ion polcntial). Specialised sense cells are
called rcccplors. Different types of sensory recepiors exist in the body.
bk>odpressure a-0rtaandcarotidartery
Th e rmorece ptors:respondtothermalstimul ation
temperaturechangeintheskin ner,eend ings
internally cells of hypothalamus
Ch e morue ptors: responding to che mical stimul ation
sensecel!sofo!factoryepithelium
2 Uselable1S.1tolist taste buds tongc;e
the various types of bk>odoxygenandwboodioxioe carotid body carotid artery
stimuli(semedata)
concentrations and pH
that originate from
cond itiomwithinthe osmotkconcentratioooftheblood osmoregulatorycentrein
human body that are hypothalamus
detectedbyp.-irtirnlar Photorece ptors:respondtoele ctrom agne tic stimulation
receptors. light
Transmission of an impulse
An impulse is transmittal 3long nen,e fibres, but it is not 3n electrical current that flows 3kmg the
'wires' d then=. Rather, the impulse is a momentary reversal in electrical potential difference in the
membrane, caused by the rapid ==ts of sodium and potassium icos into and out of the axon
The resting potential
A neurone that is not transmitting 3n impulse is said to be 31 rest. Actually, this not the case; during
the 'resting' inten"al between the transmission of impulses, the membrane of a neurone aaively
creates3ndtru1intainsanelectricalpotentialdifferencebetweentheinsideandtheoutsideofthe
fibre. The result is that, in a 'resting' neurone there is a potential difference between the inside and
the outside of the fibre of about ~70mv. (Jbe discovery d the resting potential came from the use
of an external and intem3l microdectrode on an isolated axon, the amplification of the signal and
its display on a cathode ray oseUJoscope).
Two processes together produce the resting potenti3l difference across the neurone membrane.
• There is active trnns pon of potassium ions (K"') in 3cross the membrane 3nc] d sodium ions
(Na • ) out across the membrane. The ions are transponed by a sodium---po1as.sium pump, with
transfer of energy from ATP. So potassium and sodium ions gradually concentrate on opposite
sides of the membrane. Of course, this in itself makes n o change to the potential difference
across the membrane.
• There is also facilitated diffusion of potassium ions out and sodium ions back in. The important
point here is that the membrane is far more permeable to po1as.sium ions flowing out than to
sodium ions returning. This causes the tissue fluid outside the neurone to contain many more
positive ions than are present in the cpopl3Sm inside. As a result, the inside becomes more and
more negatively charged compared with the outside,, the resting neurone is said to be polarised.
lbe difference in charge. or po1ential difference, is about - 70mV. This is known as the res ting
Figure15.5Theestablishment pote ntial. Figure 15.5 is a summary of how it is set up.
ofthe,estingpotential
316
15.1 Control and co-ordination in mammals I
The action potential
The next event in a neurone, sooner or later, is the passage of an impulse. An impulse or ac tion
pote ntial is triggered by a stimulus received at a receptor cell or sensitive nerve ending.
lntheactionpotential.theenergytransferredbythisstimuluscausesatemporaryandlocal
reversal of the resting potential. The result is that the membrane is briefly d e polarist.-d at this
point. How does this happen?
The change in potential across the membrane occurs through pores in the membrane. 1bey
are called lo n c h a nnels because they can allow ions to pass through. One type of channel is
permeable to sodium ions and anocher to potassium ions. These channels are globubr proteins that
span the entire width d the membrane. They have a central pore with a g:11e which can open and
close - they are g ated channel~. During a resting potential these channels are all closed.
The energy of the stimulus first opens the gates of the sodium c hannels in the cell surface
membrane. This allows sodium ioru to diffuse in, down their electrochemical gradient. This inOux
of sodium ioru is very rapid indeed. So the cytoplasm inside the neurone fibre quickly becomes
progressively more positive with respect to the outside. This charge reversal continues until the
potential difference ha, altered from - 70mV to+40mV. At this point, an action potential has been
createdintheneuronefibre
~ ....
1
depolarisa!ioo lonm ovementsduringthaaction
due to Na•emry potential:
1 : :•:.m:.~~ti.~::t:::t~=.ichannels for
2Na• rnanne1,oper,a ndNa•iomrush;n
(bycliflu'>K>ll)
3 lmerio<ofa,vnbecomesincreasinglymore
posiWelychargedwith respec:trotheootside
4Equally suddenly.N.1'chanlll'l,closeat the
"'""' momem as I(• cha nr,els open aod I(• ions
5 ;~::.:~\~!r.:~tarts
po<;1....,aga;n
to become k><s
Fig un,15.6Theact>onpotential
317
m (onttol and CO·Ordioation
The action potentfal then travels along the whole length of the neurone fibre. At any one point it
exists for only two thousandths d a second (2 milliseconds) , before the membrane sians to
re-establish the resting potential. So action potentfal transmission is exce,,cJingly quick - an example
o f poshfre feedback , in fact.
Almost immediately an action potential has passed, the sodium channels close and the
potassium channels open. So potassium ions can exit the cell. again down an electrochemical
gradient, into the tissue fluid outside. This causes the interior o f the neurone fibres to sian to
3 Oistinguishbetween
become less positive again. Thenthe-pot:assiumchannelsalsoclose. Finally.theresiingpotentfal
positive and negative
of - 70mV is r.,._.,stablished by the action of the sodium- pornssium pump and the process of
feedback.
facilitated diffusion.
4 Whati1themurceof
energy used to
a establishtheresting The refractory period
potential
For a brief period, following the passage of an action potential, the neurone fibre is nOl excitable.
b power an action This is the refractory period . It lasts only 5---10 mUliseconds in total. During this time, firsily there
potential7
is a large excess of sodium ions inside the neurone fibre and funher influx is impossible. As the
5 Whyisanaxonunable resting potential isprogressivelyresiored,however. itbecomesincreasinglypos.sibleforanaction
toconductanimpul1e potential to be generated again. Because of this refractory period. the maximum frequency of
immediately after an impulses is between SOO and 1000 per second .
action potential has
beenrnnducted7
The 'all or nothing' principle
Obviously siimuli are of widely different natures, such as the difference between a light touch
and the pain of a finger hit by a hammer. A stimulus must be at or above a minimum intensiry,
known as the thres hold of s timulation. in order to initiate an action potential at all . Either
a stimulus depolarises the membrane sufficiently to reverse the potential difference in the
cytoplasm (-70mV to +40mV), or it d oes not. If not. no action potential is generated. With all
sub-threshold stimuli. the influx of sodium ions is quickly reversed. and the resting potential is
However, as the interuiry of the stimulus increases the frequency at which action potentials
passalongthefibreincreases. (Note that individual actionpotentialsareallofsiandard
amplirude.) Forexample,withaverypersistentstimulus. actionpotentialspassalongafibreat
an accelerated rate, up to the maximum possible permined by the refractory period. This means
the effector (or the brain) recognises the intensiry of a stimulus from the frequenc y of action
potentials(Figurel5.7).
::;;;.:
~,,.,,,,
+""'"""'""
,Omulus
o o 0 0 0..L••L.
1
stimulus
~m
IIAAW,MO
..1.
stop, Stafls Slop,
Figure15.7Weakar,dstrnogstimuliandthettm,,l>ok1value
318
15.1 Control and co-ordination in mammals I
Speed of conduction of the action potential
We noted earlier that the presence of a myelin sheath affects the speed of transmission of the
action potential. The junctions in the sheath, known as the nodes of Ram'ier, occur at 1- 2mm
intervllls. Only at these nodes is the axon membrane =posed. Elsewhere along the fibre, the
electrical resistance of the myelin sheath prevents depolarisations. Consequently. the action
potentials are forced to jump from node to node. This is called sahatory conduc1lon (from
the Latin saltare meaning 'to leap'). This is an advantage, as it greatly speeds up the rate of
Not all neurones have myelinated fibres. In foct, non-myelinated dendrons and axons are
common ·n non-venebrate an'mals. Here, transm·s.s'on ·s normally much slower, because
the action potential flows steadily , all along the fibres. However, among non-myelinated
6 Vv'hydomyelinated fibres it is a fact that large di3meter axons trnnsmi! action potentials much more speedily
fibres conduct
than do narrow ones. Certain non-venebrntes like the squid and the eanhworm have giant
impulses faster than
fibres, which allow fast transmissions of actio n potentials (not as fast as in myelinated fibres,
non-myelinatedfibres
ofthesames.ize? however) . We saw that the original investigation of action potentials was carried out on such
giant fibres.
dirl'cliooofimpulse
d -.,- -t .. ~.:::::b
· 1· .
~· L t
~-- -
oodeofRoovier
Figure15.8Saftato,yconductf0n
Na•in
319
m (onttol and CO·Ordioation
Figure 15.9Asynapse;nsectior,
320
15.1 Control and co-ordination in mammals I
how a cholin ergic
syn apse works
structure ofca'•channe ls
inpre,synapticmembran•
(enlarged)
ea'•
\j~ 00 oo
4
=~tioool
2 Trammi!ler
mb<!anc:e
released,
3 Trill\smittersub<tance
binds,u;gge,ingermy ol
Na•ooos,aodactK>ll"°temial
inpost-synaptkmembrane
321
m (onttol and CO·Ordioation
Motor nerve endings m ake By the end of this section you should be able to:
connection1withmuscle
fibrei and trigger muscle i) describe the roles of neuromuscular junction>, transverse sy>t em tubule< and ,arcopla,mic
contraction re ticulum instimulatingcont ractioninotriat edmuscle
j) descr ib e the ultrastructu re ofst riate dmu,cle wit hpartkular refere nce tosarcome rertructure
k) explain th e slid ing filament mode l of muscular contraction includ ing the roles of troponin,
tropomyosin,cakiumionsandATP
322
15.1 Neuro-muscular junctioos and muscle crotraction I
Figure15.11The,tructu,eofskeletalmusde
Figure15.12Theutb-astructu,eofamusdefib,e
323
m (onttol and CO·Ordioation
The striped appearance of skeleral muscle is due to an interlocking arrangement of two types of
protein filaments, known respectively as thic k and thin fila m e n ts - they make up the myofibrils .
These protein filaments are aligned, giving the appearance of stripes (alternating light a nd dark
bands). This is shown in the more highly magnified electron micrograph and interpretive drawing
inFigurelS.13.
The thick filaments are made of a protein called m yosln. 1bey are about 15nm in diameter.
The longer, thin filamems are made of another pro1ein, actin . Thin filaments are about 7nm in
diameter,andareheldtogetherbytransverseOOnds,knownas Zlines . Each repeating unit of
the myofibril is, for convenience of description, referred to as a sarco m c rc . So we can think of a
myc{tbril as consisting of a series of sarcomeres attached end to end.
324
15.1 Neuro-muscular junctioos and muscle crotraction I
Figure15.14Thestrurnln,ofaneuromusrularjunchoo
a lectron
microg ra ph
microg raph
I lbe myofibril is stimulated to contract by the arrival of an action potential. This triggers release of
calcium ions from the sarcoplasmic reticulum, which surround the actin molerules. Calcium ions
now re-act with an additional prorein present (troponin) w hich, when so activated, triggers the
removal of the blocking molecule. tropomyosin. lbe binding sites are now exposed
1. Each bulbous he-ad to w hich ADP and Pi are attached (called a cha rgt...! bulbous h e ad) reacts
with a binding site on the actin molecule beside it. The phosphate group (P;) is shed at this
j lbe ADP molecule is then released from the bulbous head, and this is the trigger for the row ing
moveme nt of the head, which tilts at an angled about 45°, pushing the actin filament along. At
this step, the power s trok e, the myofibril has been shortened (contraction)
-+ Finally, a fresh molecule of ATP binds to the bulbous head. The pro(ein of the bulbous heads
includes the enzyme ATPase. which catalyses the hydrolysis of ATP. When this reaction occurs,
the ADP and inorganic phosphate (PJ formed remain attached, and the bulbous head is now
"chargt...! " aga in. The charged he-ad detaches from the binding site and straightens.
This cycle of movements is shown is Figure 15.16. The cycle is repeated many times per second
with thousands of bulbous he-ads working along each myofibril. ATP is rapidly used up, and
the muscle may shonen by about 5{m of its relaxed length. However. when the action potential
stimulation stop,, the muscle cell relaxes. Now, the filaments slide back to their original positions.
Ion pumps in the sarcoplasmic reticulum pump calcium ions back inside, and so the calcium ion
concentration surrounding the myosin filaments falls. Blockage of binding sites by tropomyosin is
restored.
326
15.1 Neuro-muscular junctioos and muscle crotraction I
Arrival of n erv11 impulH releases ea'• from the sarcoplasmk retiruh.m . ea'• iol1' cam<' remo,al
of blocking molecule fmm binding site<. Each mp;in molecule has a bulbous heitd that react,
wi!hATl' ---> ADP+P; whkhremainbound
• lhem)<lfib<i isslimu!atl'dtocootr..a
~~c!~~·r:::~"
:li=~~'.~~~7,;·m"':";.~
=i!:!~
IOith
thebkx:kir>gmolecules,remo,ir,g t hem
so the bindingsit8Sare nownposed.
• ThentheADf'moleculei<reteased
fromthern;osinhead.Thisisthe
tr'99l"f0<the·rowingmovement'ol
thehea<l,whkhtjlts byaoa'9"of
abou!45",pushingtheaelin Mament
along.Atthis<!ep, the power stroke,
them)".lfioolhasbeenshortened
(contraction)
2rl'leaseoltheADP!figge"
moYl.'lflef11ofm)'OS'n,bya
• finaly,afreshmoleculeolATl'binds
@) +- ;:;.9~~/;:=i<
-· tothem)O'inhead.Thepmte;nol!he
rnyos;nhead indudestheenzyme
AT!',,,..,whichcataly;es!hehydrnly;;.;of
AT!'. When th;.; reaaK>ll occurs, the ADP
arn1 ;00fganicphospha1e(P)fonned
remainattached . Tr,emyosrlhead;.;r.oN
'marged'again. lhecllargedhead
detachesfromthebirn!iogsiteand
maighteos..
4cyclei,repeateda! binding
shelunheralongtheac:tinmolecu!e
Swhenr.e!Vl'stimuliltKll\olmusdesceases,ca>•
ioo, retumtothes,ycoplasmicre!iculum¥Klth,e
binding5'te,becomeblodedagain
Figure15.16The,hding-filamenthypothesisofm=lecontraction
327
m (onttol and CO·Ordioation
The endocrine system i:s a By the end of this section you sho uld be able to:
slower system that controls
long· termchanges.Fertility l) explainth e rolesofthehormonesFSH,LH, oestrogen andprogesterone incontrollingchangesin
maybecontrolledbyuse the011aryanduteru.duringthe humanme nstrualcycle
of hormones . m) outline the biological basis of contracept ive pills containing oestrogen and/or progesterone
thyroid gland
Figure1S.17Tfiehumanendocrinesystem
328
15.1 The endocrine syslem I
Homxmes circulate in the blood.stream Oflly briefly because in the liver they are COfltinually being
broken down. Any bre:ikdown products no longer of use to the body are =creted in the kidneys.
So, long-acting hormone., must be secreted imo the bloodstream continuously to be effective - and
they are.
You can see immediately that hormOflal control of body funcrion is quite different from nervous
9 HOfmooe~. on secretion, control; the latter is concerned with quick. precise o:immunicatiOfl, whereas hormones mostly
are transported all work by causing specific changes in metabolism or development, often over an extended period of
overthebody.Explain time. However. these o:imrasting systems are co-ordinated by the brain. Our nervous system and
how the effects of hormOflesworkrogether.
hormones are restricted
The strucrure of endocrine glands can be contrasted with that of O!her glands in our body which
to particular cells or
deliver their secretions through tubular ducrs, such as the salivary glands in the mouth and sweat
lilrnes
glands in the skin. These ducred glands are called =ocrine glands. Their secretions pass out of the
gland via ducrs and they have altogether different roles in the body.
329
m (onttol and CO·Ordioation
By convention, the srnn of the menstrual cycle is taken as the first day of menslrnatk>n
(bleeding), which is the shedding of the endometrial lining of the uterus. lhe seeps, also
summarised in Figure 15.18 above, are as follows:
10 Explainwhatismeant
I PSH is secreted by the piruitary gland, and stimulates development of several immarure egg
by negative feedback
C()f'l/ro/.Canyouthink cells (in secondary foJHcles) in the ovary. Only one will complete development into a marure
haNpositivefeedback secondary oocyte (now in the Graafian follicle).
might differ from l lhe developing follicle then secreres oestrogen. Oestrogen is a homxne tha! has two tugets:
negativefeedbac:k7 • in the uterus oestrogen stimulates the build up of the endometrlum, which prepares the
uterus for a possible implantation of an embryo, if fenilisation takes place
• in the pituitary gland. oestrogen inhibits the funher secret:ion of FSH. This prevents the possibility
d funher follicles being stimulated to develop. It is an =ample d ncgati\"c fr'Cdback control
.i Meanwhile, the concentration of oestrogen continues to build up, increasing to a peak value
just before the mid-point of the cycle. This high and rising level of oestrogen suddenly
s timulates the secretion of LH and, to a slightly lesser extent. FSH, by the piruitary gland.
LH stimulates ovulation (the shedding of the mature secondary oocyte from the Graafian
follicle) and the secondary oocyte i, released from the ovary.
4 AsSOOflastheovarianfollicleha.sdischargeditsoocyte,Uialsostimula1estheconversiondthevacant
follicle into an additional. temporary gland, called a corpus luteum lhe capus luteum secre:es
progesteroneand,roalesscr=tent,oeslJOSerl. J>rogesteroneha.stwotargets:
• in the uterus progesterone continues the build up d the endometrium, funher preparing it for
apos.sibleimplantationofanembryo, iffenilisationtakesplace.
• in the piruitary gland progesterone inhibits funher secretion of IH. and also of FSH. This is a
secondexampleof ncg,111\'c fccdback control.
330
15.1 The endocrine syslem I
~ The levels of FSH and Ll-1 in the bloodstream now decrease rapidly. Low levels of FSH and
lli allow the corpus luteum to degenerate. A5 a consequence, the levels of progesterone and
oestrogen also fall. Soonthelevelofthesehormone.sissolowthat theextraliningoftheuterus
is no longer maintained. The c ndome trlum brea ks down and is [OSI through the vagina in the
firstfivedaysorsoofthenewcycle.
The falling k>vels of progesterone again cause the secretion of FSH by the pituitary and a new cycle
is underway. The changing levels of the hormones during the menstrual cycle and their effects on
theovariesandtheliningoftheuterusareshowinFigure15.19
levels o f
FSHandLH
in the blood
femalesu
~;:\:~-------- ... ,,,\
hormones
in the blood /
________ __________ ,/"":_....
(oolertili<atioo)
eqgcell inldlidegrowing owla tion - - ~ COf?Usluteum - ~ degenerates
1 2 J 4 s 6 7 a 9 10 11 12 n 14 15 16 11 1a 19 20 21 n 23 24 2s 26 21 2e
time(day,)
Fig..-e15.19Changinglevelsolthehormom,sduringthemerntrualcyclear>dtheireffernontheOYariesandthelinir,goltheuterus
331
m (onttol and CO·Ordioation
11 ldentifythecritical If 1hc egg is fcrtlliscd (the start of a pregnancy), then the developing embryo itself becomes
hormonal changes an endocrine gland. secreting 3 hormone that circulates in the blood 3nd m3intairu the corpus
<h.•· luteumas3nendocrinegl3ndformleastl6weeksofpregnancy.
a triggerovulation When eventually the corpus luteum does break down. the plaa,ma rakes over as 3n endo-
b causedegeneration crine gl3nd, secreting oestrogen and progesicrone. lbese honnones continue to prevent ovul3 -
of the corpus tion 3ncJ maintain the endomccrium.
luteum
For the oral contrnceptive pill to be effective, the woman must take o ne pill at the same
Figure 15.20 A commercial pack time each day for 21 d3ys. She then rnkes 3n inen pill (3 placebo - containing an iron
of"thep ill'
supplement. as a lot is lost in menstruation) for seven d3ys. or she m3y rnke no pill 31 3ll.
During this time menstru3tion occurs. This 'period" is usu3Jly lighter and shorter than is
normally experienced.
Jn the body. the hormones from the pill prevent ovulation by decreasing the release of FSH
and lli from the pituitary gland (without completely stopping their release). This restricts the
growth of follicles in the ovaries and so a secondary oocyte does not grow 3nd is not released. The
12 'Mlatdoyou endometrium still builds up in the uterus and then, later, breaks down 3gain, but menstruation is
lfilerstandbya shorterandlighterbecausethecorpusluteumhasnotgrown(intheabsenceofovulation)
'placebo'?
Nothing is without risk, but very many women who h3ve access to it find the pUJ beneficial to
thrirlifestyle.healthandrangeoflifechoices. Thelikelybenefits3ndpossiblerisksareidentified
inTablel5.2.
Table15.2Theri,ksandbenefruofthepill
• Somewomende,,.,lopnausea,headaches,tiredne" • ThereisaredtKedriskof
and mood swing, • ova f ancy,t,
• ttmaytrigge,ariseinbloodp=re.lead ingto • caoce,oftheovariesOfuterus
aninc:reasedriskofthrombosis(bloodclotsarelile· • Menstruationismoneregula, fofman)l;and
threatening pre-memtrualtemionmaybelee.sened
• The,ei,an ;naea,ed,iskofbreastcaoce,,butthisis • ThequalityofthemtKuspluginthece,vixreduces
onlytoas!fghtextent theriskofbacterialent,ytotheuterusand
the,e/o,edec:reasesthechaoceofinfection
332
15.1 The endocrine syslem I
The 'morning after' pill
lbis 'emergency' contmcqxive medication contains synthetic progesterone that works in the
lxxlyinthefallowingways:
• it delays ovulation or it inhibits ovulation if that event has not yet occurred
• it irrirntes the endomecrium so that implantation does no( occur.
Typically. the morning after pill is taken if a women has forgotten to take her combined oral
contracep1ivepillorif anO!hercomracep1ivedt,>vicehasbeenominedorusedandfailed(a
condom may have broken). It may also be used if a woman has been raped.
lnthese'mominga!ter'siruations,thesenseof'emergency'isa(quitenatural)srnteof
mind,ratherthanabiologicalfact. Thisisbecausefertilisationoccursintheoviducr,ifatall.
Afertilisedeggthenpassesdowntotheuterus, ajourneythattakes5-7days. Only then can
impbntation in the wall of the uterus occur. So there is an interval of about 72 hours in which
apregnancycanbeavoided.However, iffertilisationhastakenpbce,thenthispanicular
contracep1h measure ·s a 'chem"cal abort"on'.
~~
"
~
- A1ia(o•l"""'9Mid dlo
~!•alAmorico&
Coribbecm
-·
;
=~=Eo>1&Norm
·
}" ~
g
~:~
40 C . . - - - - -
""
'
0 +---~~~~~~~~~~~
195()...19551960-1965 1970-19751980-19851990-19952000-2005
Figure15.21Lifeexpectar,cy1950----2D05
People who make a study of human population growth (demographers) have predicted that
by 20SO the Eanh may have 10 biJHon human inhabitants - that is very many more than we
currently have. The projections suggest that the world population wm .stabilise a t this level.
Can such a total human population be susrnined by the Eanh's resources? Many people
feel that it cannor. To them, the current destruction of habitats and devastation of biological
diversiry observed on all continems is directly or indirectly due to our anemp1s to provide for
the lifestyles that =isting humans seek (or seek to maintain). It is unacceptable to them that
the environment is so abusal. To these people. one irn~iate task is to reduce the binh rate.
G:>ntraceptionhelpstomakethispos.sible.
333
m (onttol and CO·Ordioation
Plant co-ordination systems By the end of this section you sho uld be able to:
involve rapid responses as
inthecaseoftheVenus a) describetherapidresponseoftheVenusflytraptortimulationofhairsonthelobesofmodified
fly trap, but also complex l....ves and explain how the closure of the trap is achieved
interoctiom between plant b) explain th e role of auxin in elongation growth byrtimulating proton pumping to acidify cell walls
growthregulator:s.~has c) describetheroleofgibbere llininthe g e rminationofwhe atorbarley
auxinandgibberellin d) explainth eroleofgibberell ininrtemelongationincludingthe rol eof the dominantallele, Le,
Plants respond quite thatcodesforafunctioningenzymeinthegibberellinsynthesispathway,andtherecessiveallele,
differentlytodjfferent /e,thatcodesforanon-functionalenzyme
conc:entrntionsofplant
growth regulators.
334
15.1 Cont,ol and CO·Oldination in plants I
a}Theresponseofaplantstemtolight b} Lea ffallin awoody plantoftemperate climate
"""' m•ru•. .ro~S,f ofwOO<fy~O
\
fromleaftr;sue
junctionofleafbaseand
stemink,ng itud inalsectioo ageing leaf
Figure15.22Examplesofplantsensitivityandrespoo"-'<
Among the internal factors thm play a pan in plant sensitivity, the most important are the
substances ,ve call pl:im growt h n:gulalor,r and their effects. There are five major types of
compound naturally occurring in plants that we call growth regulators. The effects of these
chemicals are rather d ifferent from those of animal hormones (see Table 15.J).
Plant growth regulators:
• occur in very lo w concentrntiOfls in plant tissues, so sometimes it is hard to detennine their
precise role
• areproducedinav:irietyoftissues. rntherthanindiscreteendocrinegl:inds
• o ften interact with each other in controlling grovvth and sensitivity - some internet ro reinforce
each other's effects (synergism), others oppose e:ich other's effects (antagonism)
• if they move from their site of synthesis (and not all do), they may diffuse from cell to cell, be
acrivelytransportedorbecarriedinthephlocmorxylem
• may have profoundly differing effects. depending on whether they are presem in low or high
13 1/v'hat isthedifference
between growth and
• mayhavedifferenteffectsaccordingwhichtissuetheyareinoratdifferentstagesof
development?
developmem of a single tissue.
335
m (onttol and CO·Ordioation
Table15.)Theplant,egulato,,;:discOYe,y.,ciesandsynthesis.Thestructuralformulae
shw, how chemically d,Verse these substances are - rheydo nor require memorising
Auxins,princip11llylndoleaceticacid(IAA)
0i5"overy
lnitialfybyDaiwin.investigatingcoleoptik>sandtheircu,vatun,towardsaunilaterallightsoo,ce
Late,byothers.;ndudingthedevisingofabiok,gicalassaytofindtheconcent,atioosofauxins;n
plantorgans{s;rxetheseareatlowconcentrations) ' ~- 1- 1
rn, - cooo
• l'fomotionofextensiongrowthofstems
• Delayofleafsenescenceandleaffall
• lnhib<tionoflate,alrootinitiation
• S\\'ilch ingonofgenestopromotegem,ination
Synthesis
lntheembryosofseedsandin)'OUngleaves(except ingenetica llydwarfvarieties)
Then,areseveraltypesofg ibben,llin
Ab5"isicacid(AB.A)
Discovery
Duringinvestigatiooofbudandseeddormancyand inabscissiono ffruits
"""~.'.''rn,'"'
• Astresshom,one ~ ~ H
• Tnggeringofstomatal closurewfienleafcellsareshortofwater OH
r;,-l:~::-7- of-boo_a_ad_=_d_d_='"~"- - - - - - - - - - - - - , 0 CH, (OOH
lnmostorgansofmaturep!ants. intinyamounts
336
15.1 Control and CO·Oldination in plants I
Interactions of plant growth
regulators
Sometimes plant grovvth regulators have their effects acting on
their own, but in olher siruations, they interact with olher growth
regulators.W'henonesubstanceenhancestheeffectoftheother,
this is known as s ynergism. For example, gibberellins (GAs)
and indoleacetic add (!AA) work synergiscically in seem extension
growth.
Altematively, a growthregulatormayreducetheeffect
of anO(her. This situation is known as a ntagon ism . Some
imponantinteracrionsinvolvetheothergrowthrcgulators
(cytokinin s - theirmainroleis in cell differentiation. and
ethylene - itsmainroleis in fruit ripening).
Figure15.23TheinteractK>Oolgro.vthregu!ators
337
m (onttol and CO·Ordioation
cellwall(cellulosemicrofibrils
bo<.odt>Jotherpolysaa:harides)
' "'"'"'"' ·- j
(passiveernryabove.active
pumpingootatbaseolcell
t>jauxinelftuxpump<)
. .
.
·<_
. . .. .
. . .
.
y ;o,o;(..;thr~.· auxin-u-iggeredcel
eloogalion(acklgrawth
• REll,SEll,m;tochoodria. , h)'pO!hesis)
andGolgiapparatus)
• auxin<timulatesprotoo
•""""'
.·.· • .\ j ···. . • wal l becomesacklk
•• ....;,uote • =.,~r:!~akage of
. ..
--
. . . cellulosemkroliooland
. . . . biodiog!D'jSOCcN<kle<
• hydmly!ic:enzymesattad
expos,.'(!binding
• wall resis!ancetometchir,g
isdec,..ased - turgo<ol
••.uin-medated,..gulatioo
cellcatJSl'S<t,..tchingl
otgeneexpressioo
elongatiooofcellwal
(pfomotion/iohibitionof
geneexpr"""°"v,a
traoscriptiooregula!iool .'...;._.;:.
uaoscriptioofactors)
338
15.1 Control and CO·Oldination in plants I
The role of gibberellin in the germination
of barley
Many higher plants have been found to contain glbbcrellic add (GA). In fact, several different
forms of the molecule have been discovered and these are often referred to collectively as
gibbcrdlins. Gibberellins are formed in chloroplasts , in the embryo of seeds and in young leaves
Gibberellins are absent from dwarf varieties of pea plants, but when artificblly supplied to dwarf
varieties,theybecometall.
The highest concentrations of gibberellins occur in fruits and seeds as they are formed and in
seeds as they germinate. In the seed. gibberellins switch on genes thm promote germinmion and
go on to promote additional mitosis. thereby increasing the numbers cl cells fonned. Gibberellins
also have a role in the mobilisation of stored food in germinating barley fruits, as illustrated in
Figure 15.26, on the n=t page. During germinmion, the uptake of water activa!e.s the gene for
gibberellk acid synthesis. This gibberellic add then switches on the genes that control the synthesis
of hydrolytk enzymes. Hydrolytk enzymes catalyse the mobilisation of the food reserves. causing
the release of sugars. amino adds and fatty adds for germination and growth
4solublefoodstora tronslocallld1aembryowhete
iti,raspiredotusedtobuildnewcallsl•ugarsara
1ronslocatod1<>1hegrowiogpoimas,uaosel
frui1andseadcoa1 3gibbere0icocid1rigger,syn1he,i,
fused together
ofhydrolyricenzymes
cell,comoining,1oted a.g.omylose lstarch-malto,al
proleinlaleurooegroinsl mahase lmaltow-, glucose)
2gibbuollicociddilfusesto
prot&instora
1 g ibb&rellicocidlormedin
embryo as germination begins
339
m Cootrol 1M co-11rtlllation
the pc-.i plan!. In fact. ul! pc-.i pl ants carry at leaM: one allele (L~). which codes for a protein lh:i.1
fuoctKIIU normally in !he gibberellin--symhesis p:,thway. c:.i:olysing the formation of gibberellin.
Th:it aUele is 'domiruinf, so it may be presenl as a single allele or with two. but in either c:i.se.
gibberd.linwillbepresentandtheplan1willbetall.
Dwarf pea pi.ants carry tv.'O allele. (.le) lh:i.1 code for a non-functional procein. This enzyme fails
in the gibberellin-,iynthesis pathway. PL:i.n1s with iv.·o re<'CSSive alleles remain dwarf. lacking dr
M1pplydgibbttellinthatenablesextensiongr<1"'--iliinS1cm lengthtooocur.
"'""'"
~.::... ____]"'·
forby L•llllele l
.KtiYegibb.rellin gibberellinibsent
Flg..-a15.27Theroleofspe<ifical.....,,inthecontrolofhe,ghtmtheg.itdenpe,a,
(continued)
340
Examinationstyleqoostions I
(continued)
3 a) The steroidhormonesoestrogenandprogesteroneare
5eeretedbytheovary.Statepreciselythesitesofsecretion
ofeac:h. [21
b) Themosteffectiveoralcontraceptivesforgeneraluseare
thfl'Shold theso-calledcombinedoralcontrac:eptives(COCs},which
------- --------- ---------------
containbothoestrogenandprogesterone.
Explain how COCs produce their effects. [41
c) De1cribe two so cial implications of the use of
contraceptives 121
' [Total: 8]
(Cambridge International AS iltld A Level Biology 9700,
sii~L.tSSIJulus
Paper 04 Q2 June 2009)
' " Fig.1.1
4 a) By means of a labelled diagram show the structure
a) Stimulus B resultedinanactionpotential.Describe of a synapse [SI
whati1occurringatC, D and E [6) b) Describethesequenceofstepsbywhichanaction
b) SuggestwhystimulusAdidnotresultinanaction potentialuossesacholinergicsynapse (101
potential beingproducedwhereasstimulus B did [2) c) Outlinetherolessynapsesplayinthefunctioningnervous
[Total : 8] 1ystem [SI
(Cambridge /ntemational AS and A level Biology 9700, [Total: 20]
Paper 04 QB November 2007)
341
A Level
16 Inherited change
Ge ne tic information is tra nsmitted from ge ne ration to ge ne ration features a re inhe rited. The phe notype of organisms is determine d
tomainta inthecontinuityoflife.lnsexualreproduction.meiosis pa rtlybythe ge nestheyhaveinhe ri ted a ndpa rtlybytheeffectof
introduces ge netic variation so th at offspring resemble the ir parents the e nvironment. Genes determine how organisms develop a nd
but are not ide ntica l to the m. Genetic aosses reveal how some ge ne control in bacte ria gives us a glimpse of this process in action.
342
16.1 Passage of information from parent to offspring I
We have also seen that the nuclea r d!,ision that
The loda<e!hepos,tionsakingthern<~-
w!lerege,wsoccu,,soallele<ofthesamegene occurs during growth, kJX>wn a., mlt01<is, results
ocrupythesamelocu, intheexacrduplicationoftheseclu~=esin
the daughter nuclei formed.
ExaminatiOfl of the chromosomes at certain
gene - aspecilicleng!holtheONAof!he stagesinmitosisdisclosesanotherimportant
:~
.. -.occupyingapos;tioncalleda feature. Chromosomes occur in pairs. These pairs
of chromosomes are known a., homologou s
alle lesolagroe(aleleistheshortform p.'Urs (Figure 16.1). lbese homologous pairs of
of'al!elomorph'meanir,g'altemalive chromosomesariseinthefirstplacebecause
fo,m')
one member of the pair has come from the
maleparentandtheotherfromthefemale
parent. Homologous pairs are then maintained
bythe=actreplicationthatmkesplaceprior
to each mitO(ic division. A cell with a foll set
of chromosomes (all homologous pairs) in its
nucleusissaidtobeinthe dlploid condition .
343
m 1nheritedchange
Asexua I reproduction
Organisrn.sreproduce:ises:uallyorsexuallyandrnanyreproducebylxxhthesemechods. ln
asexual n,production a single organism produces new individuals . J\.ses:ual means "non-sexual";
no gametes are formed in :ises:ual reproduction. The cells d the new offspring are produced
by mltosL,;; (page 102), so the progeny are idemiml ro the parent and to each other. Identical
offspring produced by asexual reproducrion are known as clones .
Mammals reproduce by sexual reproduction only, but many flowering plants reproduce by
lxxhas=ualandsexualreproduction.
344
16.1 Passage of information from parent to offspring I
The process of meiosis
Once started, meio,;is proceeds Sleadily as a continuous process of nuclffir division. However, the
steps of meiosis are acplaifled in f01Jr dislincr phases ( p roph a.se, m ct:,phasc , <i n a phase and
tclo phasc). This is just for o:invcnience of analysis and description, there are no breaks between
thephasesinnucleardivisions
The behaviour of the chromosomes in the phases of meiosis is shown in Figure 16.4. For cbrity,
the drawings show a cell with a single pair of homologous chromosomes.
Meio.sis involves two divis lorui o f the nucleu s, known as m dosls I and m do,;is II . Both of
these divisions supe,jiciaff), resemble mitosis. For example, as happens in mitosis, chromosomes
are copied, appearing as sister c h rom a lkls during interphase, before meiOllis occurs. However,
early in meiosis I, h o m o logou s ch rom osom es p a ir up. By the end of meiosis I, homologous
chromoscmes have separated again, but the chrcmatids they consist of do nOl separate umU meiosis
II . So, meiosis consists of two nuclear divisions but only one replication d the chromosomes.
Meiosis I
Prophase I
What happens to chromosomes during prophase I is quite ccmplex and difficult to see when you are
looking at dividing cells under the microscope. However, you can follow the changes in Figure 16.5.
The chromosomes appear under the light microscope as single threads with many tiny
bead-like thickenings along their lengths. These thickenings represent an early stage in the
processdshorteningandthickeningbycoilingthatcontinuesthroughoutprophase.lnfact,each
chromosome has already replicated and consisls d two chromatids but the individual chromatids
arenotvisibleasyet.
Crossing over
Within the homologous pair, during the coiling and shortening proces..s, breakages d the
chromatids occur frequently. Breakages occur in non-sister chromatids at the same points along the
length of each. Broken eritls rejoin more or Jess immediately. but where the rejoining is between
non-sister chromatids. swapping d pieces d the chromatids has occurred. hence the name
'crossing over'. This results in new ccmbinations d genes on the chromosomes.
lbe point of the join between different chromatids is called a
c hia.sm ., (plum/: c h lasm ata). Vinually every pair of homologous
chromosomes forms at least one chiasma at this time, and two or more
chiasmata in the same homologous pair is very common.
lnthelaterstageofprophaseltheattractionandtightpairing
of the homologous chromosomes ends, but the anracrions between
sister chromatids remains for the moment. This attraction of sister
chromatids keeps the homologous pairs together. The chromatids are
lnlateprophaselthetwocentriolesdthecentrosomepresentin
animal cells (page 19) duplicate. The two centromeres then scan to move
apanasapreluderospindleformation.Plamcdlsdonothavecentrioles
Figure 16.4 f'hotomKJographolhomok,goll5ch,~= Finally,thedisappearancedthenudeoliand 11uclcarc1welopc
f>eldtogetf>erbychiasmata mark.stheendofprophasel.
345
m 1nheritedchange
prophasa l(late)
~~~~
toconst1tolchromancisSi1eswhere
chmmalldshawbrokenandrejoif>ed.
ch""""1ta "9"""'""'""tileas
anaphase l
-·"'~-"-'P",ate
'""""'"'=-arepYlled
~ : : ~ ~ : • J)Olesol !hesp<lde,
Figure 16.STheprocessofmeia.is
346
16.1 Passage of information from parent to offspring I
1 Homologoosch~- SOncepairsofoomologou,
rn~-llave,epa,ated.
~:::=~~~oa~{hoden thesitesofaos:singOYer
by coiling arenotapparem
Metaphase I
Next. the spindle forms, and the homologous pairs become atrnched to individual microrubules
of the spindle by their centromeres. The homologous pairs are now arranged at the equatorial
plate of the spindle framework - we say that they line up at the centre of the cell. By the end
of mernphase I, the members of the homologous pairs smn to repel each other and separate.
However, at this point they are held together by one or more chiasmata and this gives a temporary
butunusualshapetothehomologouspair.
The cl,--
Anaphase I
of each homologous pair now move to opposite poles of the spindle, but with the
individualchromatidsremainingattachedbytheircemromere:s.Theanractionofsisrerchromatidshas
lapsed.andtheyseparateslighdy - botharedearlyvisible.However,theydonotseparateyet;theygo
rothesamepole. Con.sequendy,meiosislseparateshomologouspair:sofch--butnotthesister
chrcmatid:sofwhid,eachiscomposed.
Telophase I
The arrival of homologous chromosomes at opposite poles signals the end of meiosis I. The
chromosomes tend to uncoil ro sane extent, and a nuclear envelope re-forms around lxxh nuclei.
The spindle breaks down, but these two cells do not go into inrerphase , rather they continue into
meiosis n. which takes place at right angles to meiosis I. There may, however, be a delay between
meiosislandmeiasisll, asin~enesis(pageJ44)
2 'NhatistheJTliljorc:ornequenceoftherebeingnointerphasebetweenmeiosislandmeiosisll7
3 Atprroselywhichpointsintheprocessofmeiosis{Figure16.S)dothefollov..ingeventsoccur?
a thenuclearenvelopebreabdownandthespindleform1
b homologouschromosomespairup
c thepresenceofchiasmatabecomesvisible
d chromatids separate at their centromeres
e homologous chromomme1 separate
f haploidnucleiform
347
m 1nheritedchange
Meiosis II
Meiosisllisremark3blysimilartomitosis:
• Prophase ll
lbe nuclear envelopes break down again, and the chromosomes shorten and re-thicken by
coiling. Cerurioles, if present, duplicate and move ro opposite poles of the cell. By the end of
prophasellthespindleappararushasre-formal. butmrightanglestotheoriginalspindle
lbe chromosomes line up at the equator of the spindle, attached by their centromeres.
• Anaphascll
lbecentromeresdivide andthechromatids moverooppositepolesofthespindle, centromercsfirst.
• Telophasell
Nuclear envelopes form around the four groups of chromatids, so that four nuclei are formed
Now there are four cells, each with half the chromooome number of the original parent cell.
Finally, the chromatids - now recognised as chromooomes. uncoil and become apparently
Allele:ooeoftlvoormore dispersed as chromatin. Nucleoli re-form.
altemiltivenudeotide The process of meiosis is now complete, and is followed by division of the cytoplasm
sequences at a single (cytokinesis,pagelOI).
geoekx:U5,soallelesare
variantfc.-msofagene
Forexample,thealleles
oftheABObloodgroup
Meiosis and genetic variation
geoearefoundatakx:us There are differences in the generic information carried by different gametes. This V3riation arises in
on chromosome 9, 1\ith meiosis and is highly significant for the organism, as we shall see. lbe reason why the four haploid
theallelesindudingt',(' cells produced by meiosis differ generically from each O!her is because of two important features of
andl°.Oiploidbodycells
containtwocopiesofeach
homologoU5 chromosome, • the indepcndc m assort ment of mat er nal a nd paternal homo logou s c hro m oe;.omes. lbe
sohavetwocopiesof way the chromosomes of each homologous pair line up at the equmor of the spindle in meiosis I
chromosome 9, and so have is entirely random. Which chromosome of a given pair goes ro which pole is unaffected by
twocopiesofthegene (independent of) the behaviour of the chromosomes in other pairs. Figure 16.7 shows a parent
Thesemaybethe!dlle cell with only four chromosomes, for clarity. Of course, the more homologous pairs there
allele{horrozygous),for areinthenucleus,thegreaterthevariationthat ispossible. lnhumansthereare2Jpairsof
examplet't',or f' florl" chromosomes, so the number of possible combinations of chromosomes that can be formed a, a
J",ortheym;iybedifferent result of independent assortment is 2'13 • This is over 8 million.
alleles(heterozygous),for
• the crossing over of scg,nc nts of individual maternal and pmcr nal homologous
examplel" f',cxl"l°cxf'I°
c hro mosom es. This re,ults in new combinations of genes on the chromosomes of the haploid
The gene for prowcing the
cellsproducal, asiJJustratalin Figure16.8. Crossingovergeneratesthepossibilityofanalmost
haemogobin !3-polypeptide
hasanumberofalleles unimaginable degr= of V3riation. For example, if we were to assume for sake of discussion that
Twoofthesearethenormal there are 30000 individual genes on the human chromruome complement. all with at least two
allele,Hb",andthesicklecell altemativealleles,andthatcrossingoverisequallylikelybetweenallofthesegenes.thenthere
allele,Hb',givingHb"Hb" are 2l()OOO different combinations! Of course, all these assump(ions will be inaccurate to varying
and Hb' Hb' as homozygous =tents. but the point that virtually unlimited recombinations are possible is established .
geootypesandHb"Hb'asa
heterozygous genotype. Finally, funher genetic variation in the progeny of organisms that reproduce sexually is assured by
the random fusion of male and female gametes in sexual reproduction.
A note on 'recombinants'
Offspring with combinations ofcharacteristicsdifferentfrcmthoseoftheirparentsarecalled
recombinant.~. Recombination in genetics is the re-assortment of alleles or characters into different
combinations frcm those of the parents. 'We have seen that recombination occurs for genes located
on separate chromosomes (unlinked genes) by chromosome assonment in meiosis (Figure 16.7), and
forgcnesonthesamechromosomes0inkedgenes)bycrossingoverduringmeiosis(Figure16.8).
4 What are the essential We shall see in the next section, how crossing over, random assortment of homologous
differences between chromosomes during meiosis, and random fusion of gametes at fertilisation may also lead to the
mitosis and meiosis?
expression of rare, recessive alleles, such as those that are re,ponsible for albinism, sickle cell
anaemia. haemophiHa and Huntington"s disease.
348
16.1 Passage of information from parent to offspring I
ln meios isl !hepairsoirtomologous
rn,~-moYe!Olr,epqua!O<
ol!hespind~and ineup,anoomly
Then,whenhomologoosch~-
"-'P'""•tetheymoYl'to tnl'nearestpoie
Thismayocc,.
,__,, __ __ _
Th<etteruofgen<tic--,ooo
att5hown in <>n<J),Wof
=~~m;:=.:.
,,Hoo-doslou>
<hromatidsofeach!>omoloc.,oo,
-.-------------
po~ atpro!)fl.,.,I
'Mlffithediromatid>br~
atcorJMpOnd"'!lpoolt>
olonsilh<~ le"lllh, II,« -------.,
• ---
. . . . . .Cl. . . . . . . . . .~ ~ ·=:,~"""
Thediromatidsfiooly
....,.,.,.000,,..,...,to
l>aploidnoclNinm<iosi,I,
~=-."'!:"'genetic
Figun,16.BGenetKvariationduetoaossiregOYerbetweennoo·siste,dirnmatids
349
m 1nheritedchange
Patterns of inheritance are By the end of this section you should be able to :
explainedbyusinggenetic
a) expla inthe termsge ne,locus,alle le ,dominant,reces,ive,codominant,linkage,tfflcro",F1and
diagrams.The results
F2,phenotype,ge notype ,homozygousandheterozygou.
ofgeneticuossesare
analysed statistically using b) u,e gene ticdiagram,tosolve probl e m,involvingmonohybridanddihybridcrosses, includi ng
thedii-squaredtest thoseinvolvingautosomallinkage,,exlinkage,codomina nce,multipleall e lesandge ne
c) u,egeneticdiagram,tosolve problem,involvingtestcrosses
d) use t he chi-square dtentotenthe ,ignificance ofdiffe re ncesbetw""nobservedand expected
35-0
16.2lherolesofgenes indeterminingthephenotype .
5 At the time of
Mendel's experiments
it was thought that
the characteristics of
parents'blended'in
theiroffspring.What
features of Mendel's
methods enabled him
dwarfv.tallplants
to avoid this error? Figu re 16.10 Featuresoftheganlenpea
351
m 1nheritedchange
~--
pure.brH dinglallplants
pea,;lrom
pu .... breedingdwarfpl ants
pan1ntal
g..-.aration(P)
sell-feniis.nion
was,,wmro
j
m;ny.-d•produa,d
at!hk sta~
flowe-ed;r.d;,jlowedto
..11.fe rtilile
j
seeds produced
(Mendo,fllCOtded1064seeds)-
pi;r,ted;r,dgOM>.;r.dfo....-.d
genaration(F,l tobeinthe,atioof
Jtall:1dwarlplanH
(MendejfflCOfded787t;,jl :277dw;irl,ratioof2.S4 : 1)
Figure16.11Thestepsofthemonohyt,ridcross
In the monohybrid cross it appears that the 'dwarr characteriS!ic disappeared in the first generation
Dominant:anallele\\itha (F1 generation) but then reappeared in the F, generation. We can explain what is going on if there
phenotype that is expressed is a factor controlling 'dwarfness'. This 'factor' remained intact from one generation to another.
evenwhenpresentwith However, it does not have any effea (we say it does not '=press itselr) in the presence of a
anallelethatisrecessive similar factor for 'tall'. Jn other words, 'tall" is a dominant characteristic and 'dwarf is a n,ccs;;i\'e
toit.Forexample,inthe characteriS!ic. So there must be two independem factors for height. one from one parent and the
ABObloodgroupgene,I"
otherfactorfromtheotherparent.Asexcell(gamete)mustcontainonlyoneofthesefactors
isdominanttol".Therefore
Today we see that these 'factors' are alleles of the gene for height in the pea plant. When
apersonwiththegenotype
l" l"hasbloodgroupA the tall plants of the F, generation were allowed to self-fenilise, the resulting members of the F,
because only the dominant generation were in the ratio of 3 tall plants to I dwarf plant. The genetic diagram in Figure 16.12 is
allele is expressed the way we explain what is going on.
Fromthemonohybridcrossweseethat:
Recessive:anallelewith e withinanorganismaregenes(called'facrors'whenfirstdi=ered)controllingcharacteris!ics
aphenotypethatisnot
expressed when an allele
thatisdominanttoitis • therearetwoallelesofthegeneineachcell
present.Forexample, e oneallelehascc.nefromeachparern
1°isrecessivetol",50a e eachallelehasanequalchanceofbeingpa.ssedontoanoffspring
per50nv.iththegenotype • the allele for tall is an a lternative fom, of the allele for dwarf
l"l"hasbloodgroupA, e theallelefortallisdominantovertheallelefordwarf.
andaper'iOOcanonlybe We call this the Law of Sc,gregadon.
bloodgroupOiftheyare
homozygousrecessive,1"1°. Duringmeiruis,pairsofallelesseparntesothateachcellha.sonealleleofapair.
Allelepairsseparateindependentlyduringthefonnationofgametes.
352
16.2lherolesofgenes indeterminingthephenotype .
Phenotype:thephysical,
detectable expression of
,,..M,,,,.,
par<tntal(P)
--
a small m.mber of alleles offspring (f 1)
of a particular gene, it may
begeneticallydetem1ined eachheterozygou,
{e.g.humaneyecolour}, parempmduc:eslW<I
giving rise to discontinuous kinds o f~tes'M!h
I \ --~=~h
variation. When the ,espectrthe T allele
phenotypei:scootrolledby
theadditiveeffectsofmany seg,egationomn
geoes{polygenic},itmaybe
affected by the environment
F, .. lfed
!'~.=~d
aswella1genes(e.g
humanheight),gMngriseto (mebsis) move!oOJ)!'OS'te
continuous variation / ",.. 1D'-'5ofthespinlle
0J 0. ·-·-
Genotype: thepartic:ular
allelesofageneatthe
~ s e p a < ated imo
appropriate locus on both different9ame1e<
copies of the homologous (=t.twoiSegegation)
chromosomes of its cells
{forexample,IAl").ltis
50metimesdescribedasthe
genetic constitution of an
this is called a Punnett grid-
organismwithrespecttoa it<hows allthe possible
pmgenyofthecros,
114n
heterozygous
hefebothdaod'(gametes
tal l "'ants
possiblegeootypes
ca nbeT0<t, gi,,;n9four
Figure 16.12 Genetic diagram showing the behaVK>\l, of alleles ;n the mono hybrid cross
353
m 1nheritedchange
Ho mozygo us:aterm
_,_parenta l
Mter:'gou<
~T
describing a diploid
organismthathilsthesame
alleleofageneatthegene'1 (""-'OSis) (meiostl)
loru1onbothcopiesofthe
homologoo1chromosomes
initscell1{e.g.HbAHb"}and
therefore produces gametes
with identical genotypes
0 b 0 b
{all Hb"}. A homozygote
isanorgani!.mthatis
homozygous.
Heterozygo us:aterm
demibingadiploid
organism that has different
allelesofageneatthe
gene'slocu1oneachofthe
homologous chromosomes
initscells{e.g.HbAHb'} Thefractions,epr=t
and therefore produces tMprobabil ilil's!hat
gametes with two different partic:ulargamete-,and
genotypes{Y.!Hb"and
Y.,Hb') . Aheterozygote
!ii.:~;1'""iype,)
isanorganismthatis
--
heterozygous
off, pring (F,)
pl>eootype-,ralio
Figure16.13Geneticdiagramsl,.c,rnngthebehav,0Urofallelesinthemonoh){lridtestaoss
354
16.2lherolesofgenes indeterminingthephenotype .
Codo minant:allelesthat
Modification of the 3: 1 ratio
arebothexpressedifthey Jn certain types ofmonohybrid cross the J: 1 ratio is not obrnined. Two of these situations are
arepre5enltogetherina nowiJJustrated
heterozygousorgani1111. For
example,all~esl"andl' of
theABObloodgroupgene Codominance -when both alleles are expressed
arecodominant.Therefore, Jn the case of some genes, bol:h alleles may be expressed, rather than one being dominant and the
in a heterozygous person, t" oiher recessive in the phenotype, as has been the case up to now. When bol:h alleles are expressed
I", both alleles areexpre;sed codominantly we know that both are transcribed into messenger RNA.
andthebloodgroupis An example of codominance is the case of the haemoglobin 13-polypeptide gene, where the
AB.lnthecaseofthe
phenotype of a person who has the genotype Hb" Hb" is unaffected by sickle cell disorder. the
haemoglobin a-polypeptide
phenotype of a person who has the genotype Hlr' Hb' is the less severe sickle cell trait and
gene, codominance means
thatthephenotypeofa the phenotype of a person who has the genotype Hb' Hb' is the more severe sickle cdl anaemia.
personwhohasHbAHIJ"' You can see that, in a cross between parents both of whom have the sickle cell trait. the probability
isunaffectedbysid:lecell tha1anyoffspringwillhavesicklecellanaerniaisoneinfour(Figurel6.t4).
disorder,thephenotypeof
apersonwhohasHbAHb'
isthelessseveresic:klec~I parental
traitandthephenotypeof
apersonwhohasHb'Hb'
pheoo!W" ~::=:~. t ~ : ; ; : ~..!
genotypes Htr' Hb' Htr' Hb'
isthemoreseveresicklec~I
(meiosis) (meiosis)
/ '\ /'\
8 8 8 8
7 Constructforyours~f
{using pencil and
paper) a mooohybrid
cross between cattle of
avarietywithageoe
for coat colour with
codominantalleles,
'red'and'white'
coat.Homozygous
parentsproduce'roan' offspring (f1 )
offspring{redand
white hairs together) Hb' Hb'
Predict what offspring
youexpectandinwhat
proportiomwhena
siblingcross(equivalent
to'selfing'inplants)
occurs between roan
offspring
Figure 16.14 Ge<>etic diagtam shO\,ing the inheritance of sickle cell fiaemoglobin an example of rndom,nance
355
m 1nheritedchange
Table16.1TheABOblood
g,oups-pfieootypPSand
genotypPS
r·\
l" f ' /
B
f' isrl'Cl'SSM'
to l "
·· 0 ·· 0
,y-.lt
,l'
,.•
lt4 1• 1°
lhefractions,epr'-"<'nl
the probabil ities that
parti::ulargarnetl'Sand
:kcJr"'lypl's)
offsp ring
--
geootypesra!KI
groopA ....
Figure 16.15 Genetic diag,am shm,ing the inh!'fitance of human blood ABO group alk>les
groopO
356
16.2lherolesofgenes indeterminingthephenotype .
Sex chromosomes
Jn humans, inheritance of sex is controlled by specific chromosomes known as the sex
chromosomes. Each of us ruts one pair of sex c h rom .-.m es (either XX or XY chromosomes)
along with the 22 O!her pairs. (Ihese cxhers are known as a m,- .mal c h rom osom es.)
Female gametes produced by meiosis all carry an X chromosome, but half of male gametes carry
an X chromosome and half carry a Y chromosome. At fenilisation, a female gamete may fuse with
a male gamete carrying an X chrcmoscme, which results in a female dfspring. Alternatively, the
female gamete may fuse with a male gamete carrying a Y chromosome, which results in a male
offspring.
So, the sex of offspring in humans (and all mammals) is decided by which type of male gamete
fuses with the female gamete - that is, by the male.
Note also that we would =pect equal numbers of male and female dfspring to be produced by
abr"""1ingpopulationovertime.
357
m 1nheritedchange
--
pa,..,,,tal(P)
,,G)/
~
",,0
(meiosis)
humanfl'male(9)
,, G)/
"
(meiosi'.I)
"
,, G)
't,XY
Dff'Jlring(F1 )
Figure16 .16XandYc:hromo,omesandthedeterminabcrncfsex
this hormone at this stage is to inhibit the devdopment of female sex organs and to cause the
formation of the testes. scroium and penis. In the absence of a Y chromosome a n ovary develops.
Then. panJy under the inOueoce of hormones from the ovary, the female reproductive structures
develop.
Sex linkage
It is obvious thm a ny gene present on the sex chromosomes is likely to be inherited with the
sex of the individual. These genes are said to determine SCJ<. linked c haractcri~tks. However.
theinheritanceofthesesex-linkedgenesisdifferentfromtheinheritanceofgenesontheOl:her
drro-=s present. This is because the X chromosome is much longer than the Y chromosome,
so many of the genes on the X chromosome are absem from the Y chromosome. Jn a male (XY),
an allele present on the X chromosome is most likely to be on its own and will be appc,ren! in the
phenocypec,·cnifiti,;;a rcccsslvcallclc.
Meanwhile, in a female, a single recessive gene is often masked by a dominant allele on the
other X chromosome, and in these cases the recessive allele is not expressed. A human female can
be homozygous or heterozygous with respect to sex-linked characteristics, whereas males will have
only one allele.
'Carri ers'
lnaheterozygousindividualwithonedominamalleleandonerecessivealleleofagene,the
recessive allde will not have an effect on their phenotype. The individual is known as a carrier.
They 'carry' the allele but it is not =pressed. h is masked by the presence of the dominam allele.
Consequently, female carriers are heterozygous for sex-linked recessive characteristics. In the
case of a male (XY) , the u npaired alleles of the X chromosome are all =pressed. The alleles on the
short Y chromosome are mostly o:incemed with male structures and male functions
358
16.2lherolesofgenes indeterminingthephenotype .
However, there are some recessive genetically-inherited conditions caused by recessive alleles
on the X chromosome. Examples of these are Duchene muscular dy.s1rophy, red-green colour
blindness and haemophilia. NOl:ice that these examples are gcnctkally-inhc rltc d conditio ns.
The consequence of the presence of one of these alleles may be quite different for males and
females. lbis is because, if a single recessive allele is present in a male human, the allele will be
expressed. Meanwhile, a female must be homozygous recessive for a sex-linked characteristic for
the allele ro be =pressed . A female who is heterozygous for one of these will be a 'carrier'. We can
illustrate this point by looking at the inheritance of the conditions of red-green o:ilour blindness
and of haemophilia .
Colourblindne ss is d et ectedbymulticolouredtest
parental (P)
cuds.Amosa ic ofdots isarra ng edonth ecards so
thatthose withnorma l colour v ision seea pa ttern pheoot;pes carrier female(9) oormalmale(())
tha tis notvis ible tothose with colourblindne ss.
xBxb ,,,
,",0
(meiosis)
,",©
(meiosis)
,,© ,/ ,,,@ ,/
11, xBy
--
offspring (F1)
,,,
00<mal
cdou, visioo cokuvisioo
'-----v----'
femate. m,.,
19) 10>
Figure16.17Thedetectionandinhelitanceolre<kjreencolourblindne,.s
359
m 1nheritedchange
Table16.3£xpectedratioofofuprir,gfrnmadihybridrnru
plancr9'™"'from
~o ~o
homozygous homozygous
roond.yellow wrinlded.green
allm,md.
yelklwpeas
0
(selled)
,,.,.
wriril<Jed
0 0 0 0
Figure16.18Adihybridcross
It wa.s nOl:iced that two new combination.s ( rccomblnalion....), not represented in the parents,
appeared in the progeny: either "round" or 'wrinkled" seed.s can occur with either 'green' or 'yellow'
cocyledorui.
Thu.s the two pair.s of facrors were inherited independently. That means either one of a pair of
contrasting ch.aracrers could be passed to the next generation. This tells us that a heterozygous
plant must produce four types of g:,metes in equal numbers. This is expbined in Figure 16.19.
From the dihybrid cross we can conclude a l.aw of Independent Assortment :
360
16.2lherolesofgenes indeterminingthephenotype .
----
parental (P)
nomozygou,
rooodandyelow """"''"""'
wrinkled and green
""""=
,.____ the alleles tor 'rO<.Od' and 'yellow' aft'
he!erozygous
,oundandyeloN domooant;thoef 1 p<ogenyareall
rnundyelklwpeas
F1 se lfed
"" (melO'ilS) assortmemoa:u1S oe
~ ~ ~:::.~::::::it:::
8808~ ~""'""'"'"'"""'"'~~~
"' / Y'Y (bothrfand'i')are foonedmeqoal
propomoo,(1ndependen1
a'5ortment)
\.
. _I___ _
+ - - - randomlenilisationbetweeniOl fou r
game!espmdua-;9 : 3 : 3 : 1 ratiool
offspring (F,l
1 RRyy
2 RrYY
2 RRYy
2 Rryy 2 rrYy "'"
4 RrYy
- ~,.,.
roond
361
m 1nheritedchange
--
were tall and
cut-leaved.The5e\\'ere
~bsequentlycro,sed parental (P) plan::,z,,ro.vn
and the progeny of homozygous hetemzygoos
this second generation wrink~l"'•<wilh murnlpeaswith
were in the ratio greenrotyledom )"'liowrotyledon,
• talland RrYy
cut-leaved 9
(meiosis)
• talland
IX)lato-leaved 3 ,,//~
• dwarfand
cut-leaved
• dwarfand
3 8888
IX)lato-leaved 1
Draw a fully
annotated genetic
diagram to show
genotypes of the
parents and progeny.
12 lnabreedofax:ker
~nielblackcoat
{B allele}isdominant
toredco;it{b allele)
and!iOl idpattern
{S allele} isdominant
to spotted pattern
(sallele). offspring
a Whatarethe
-- ""'
1XJ11iblegenotype1
""' "''
..
of a spaniel with
geooiy"°"ratio
a solid-patterned
black coat?
b Howcouldyoufind ,.=•• ,..,.
wrink~
362
16.2lherolesofgenes indeterminingthephenotype .
Introducing Drosophila
lhe fruit fly (Drosvpbila) commonly oa:urs 3round rotting vegetable materi:il. lhe most ccmmon form
is dten mlled the 'wild type', to differentLue it from the v:irious n3turaUy occurring mumm forms.
Drosophila has become 3 useful experimental 3nimal in the study of genecks because:
e Drosophi/ah3sfourp3irsofchromosomes(Figure16.21)
• from mating to emergence d 3dult flies (generation time) only rnkes 3bout 14 weeks
e onefem3leproduceshundredsdoffspring
wild type Drosophila • fruit flies 3re relatively easily h.andled. cultured on sterilised artificfal medium in gl3ss bottles 3nc]
theyc.mbetemporarily3n:iesthe1isedforsettingupcultures3ndsoningprogeny.
A dihybrid cross can be shown in Drosophila, for a:3mple by crossing nom,3l flies (wild type)
with flies homozygous for vestigfal wings 3nd ebony body (Figure 16.22). These characteristics 3re
controlled by genes tit.at 3re nor on the sex chromosomes . The genes concerned in this Drosophila
Druophilawith l>moph/lawith
n<lffl"liMwing,andoormal bOdy ""'tigial'Mr,g,ar>debooybody
(wildtwel (JOO!a ntform)
13 Oef1newhatismeant
by the term mutant
organism{orcell}
14 a Construct a genetic
diagram for the
dihybrid UOS! shown
inFigure16.22,
using the layout
91venin
Figure16.19
b Determinethe
genotypes of the
offspringoftheF,
generation
c ldentifywhich
of these are
recombinants
oormalwing,
oormal body
364
16.2lherolesofgenes indeterminingthephenotype .
Firstwecakulatex'(Inthis example,x'is0.47)
We now o:insult a rnble of the dlstribmion of)( to find the probabilhy of obrnining a deviation
by chance alone as large as (or larger than) the one we have
The table takes accoum of the number of independent o:imparisOfls involved in our test. Here
there were four categories and therefore there were three comparisons made - we call this 'three
d egrccsoffrccdom (d.f.)'.
Table16.6Tableofx_1 distribution
Degrees of ,_,_,ob_.._iW~cy.,_o•_m_~ '-~-'",' - - ~ - ~ - - ~ - ~ ~ - - ,
freedom
The probability given by the table suggests that the differences between the results we expected
and the outcome we observed is due to chance.
15 lnthedihybrid • If the value of)( is bigger than the critical value highlighted (a probability of 0.05) then we can
te1tcro11between be at least 95 per cent o:infidem that the difference between the observed and expected results is
homozygomdwarf significant.
pea plants with • lfthevalueof)(issmallc r thanthecriticalvaluehighlighted(aprobabilityofO.OS)thenwecan
terminalflowers(ttaa) beconfidemthatthedifferencesbetweentheobservedandexpectedresultsareduetochance
andheterozygomtall Jn biological experiments we rnke a prob.~bility of O.O S or larger to indicate that the differences
pea plants with axial
between observed (0) and expected (E) results are n o t significa nt. We can say they are due to
flowers{TtAa},the
chance.
progeny were
• tallwithaxial Jn this example, the value (0.47) lies between a probability of 0.95 and 0.9(). This me-ans that a
flowers: 55 deviationofthissizecanbeexpected90-95percemofthetimestheexperimentiscarriedout.
• tallwithterminal There is clearly no significant de\."iation between the observed (0) and the expected (E) results.
flowers: 51 The chi-squared test is similarly applicable to the results of test crosses and monohybrid crosses.
However, in any chi-squared lest that does not confirm that the results confonn to the anticipated
• dwarfwithaxial
flowers: 49 result (i.e. if it gives us a value for )( that is bigger than the critical value meaning that the
probability of getting that result is s maller than O.OS), then we must reconsider our explanation
• dwarfwithterminal
of what is occurring. In this outcome. the statistical test gives no clue as to the actual position or
flowers: 53
behaviorofthealleleso:incemed. Furthergeneticinvestigationsarerequired.
Usethe)Ctestto
determine whether
ornotthediffereoc:e
between these
observed results and
the expected results is
significant.
365
m 1nheritedchange
Queen
Ahoneybeecommmitytypkal~comi'.lts
ol20000----80000 indivK!uals.olwhich
• surve~r,gf0<f"'-'Or,gsitl'S
o gua,ddutya!hiveer,trar.c:e
• f0<agir,gf0<foodandwa1e,
· :;ic~~~boo~":s
Figure16 .23Honeybees-twogenotypesbutthreephenotwes
366
16.2 How mutations may affect the phenocype- studies in human genetics I
16.2 continued ... How mutations may
affect the phenotype - studies in
human genetics
Studies of human genetic By the end of this section you should be able to:
conditions have revealed
the links between e) explain thatgene mutation occursby,ubstitution,de letionandin,ertionofbue pairsinDNA
genes,enzymesandthe andoutline howsuchmutationsmayaffectthe phenotype
phenotype f) outline theeffe ctsofmutantalle lesonthe phe notypeinthefollowinghumanconditions:
a lbinism,skkle ce llan ae mi a,hae mophiliaandHuntington'sdi,ease
g) explainth e re lationshipbetweenge nes, e nzymesandphenotype withrespecttothe genefor
tyrosinase thatisinvolvedwiththe productionofme lanin
Table16.7Differentwa"jSthebasesequeoce;nagenemaybealtered
367
m 1nheritedchange
Wehavealreadynotedthepotentialadvantageinhavingthesicklecelltrait.Malariaisthe
most important of all insect- borne diseases (see Figure 10.7, page 197). The malarial pathogen
16 Explain how the muse Plasmodium COlllpletes its lifecycle in red blood cells. but it cannot do so in red blood cells
of sickle cell trait and containing the abnormal form of haemoglobin (lib'). People with sickle cell trait are protected to a
sickle cell anaemia significant =tent. Where malaria is endemic in Africa, possession of one mutant allele (i.e. having
supports the concept sickle cell trait but not sickle cell a>1aemia) is advantageous.
of a gene as a linear Tum back to Figure 10.9 ( page 199) now ,md "ale the correlmions between the distributions ef
sequence of ba= the sickle cell allele and ofthedistn"b111io>1 of ma/anti in Africa
368
16.2 How mutations may affect the phenocype- studies in human genetics I
,.,
o-}
ofemak, uruffected
o-}
oh-male affec:!ed
b<fthsequenceol
offspring(s,~ings)
(coosin<)=mmanguinity
T,mothy
Figi.-e16.24Anexampleofahumanpedigieec:hart
17 lnthehumanpedigreechartinFigure 16.24:
a whoarethefemalegrandchildrenofRichardandJudith
b who are Alan\
i grandparents
ii uncles
c how many peoJE in the chart have parents unknown to us
d whatarethenamesoftwooffspringwhoarecousins?
18 Explain why conditiom controlled by a mutant, dominant allele are likely to appear in every
subsequentgeneration,butcharilcleri1ticsduetoasinglerecessiveallelearelikelytoappear
infrequentlyinthefamilypedigreechart
Wecanu..-apedigreecharttodecectconditionslikelytobeduetodornin3ntandrecessivealleles.
Jn the case d a characteristic due to a dominant allele, the characteristic tends to occur in one or
more members of the family in every generation. On the oiher hand, a recessh "C charaaerisck is
seeninfrequently. oftenskippingmanygenerations.
Albinism
Albinism is a rare inherited condition of humans (and other manunals) in which the individual has
a block in the biochemical pathway by which the pigmeru melanin is fanned. Albinos have white
hair, very light coloured skin a nd pink eyes. Albinism shows a pattern of recessive monohybrid
inheritance in humans (Figure 16.25). In the chart shown of a family with albino members, albinism
occurs infrequently, skipping two generations altogether.
369
m 1nheritedchange
,.,
Do)""""'
0~ D)...,
;;""'°"' 0 a
Albinopeoplemus1be
homoz)'90IISf0<the
rece<sNeallele(pp)
Peoplewj!hoonnalsl<in
pigmenta1""1maybe
"°"""'"'""""'
(PP)o,carrie!S(Pp)
i!.!'lllllllllllllll~
Thi,isaiypk;altamotyueefor'1hefiranceofa
characteri<1iccontrolledbyarecessivealele
Huntington's disease
Huntington"s disease (HD) is due to an autosomal d o mina nt allele on chromosome 4 . The
mutation takes the form of repeats of three nucleotides (CAG) between 36 and 120 times at a
particular locus in the gene. The function of the gene, or rather of the protein it codes for, is
unclear. but it appears to be active in the brain.
Thediseaseisextremelyrare(I caseper20000livcbirths). Affectcdindividualsarealmost
cenainly heterozygous for the mutant (defective) gene. Appcarance of the sympwms is usually
delayeduntilthcageof40--S0years,bywhichtirnetheaffectedperson - unawareofthepresence
of the disease - may have passed a copy of the dominant allele to one or more of his or her
children. Any child of an affected persor, has a SO per cent chance of inheriting the o:indition.
The disease takes the form o f progressive mental deterioration. which is accompanied by
involuntary muscle movements (twisting, grimacing and staring in •fear'). A person with HD then
loses all control of his or her mental and physical abilities, and death occurs within 10 years. The
disease is named after the American d0<1or who first investigated the o:indition. There is no known
370
16.2 How mutations may affect the phenocype- studies in human genetics I
Haemophilia
Jn the circulatory system of a mammal , if an injury occurs there is a risk of uncOfltrolled bleeding.
This is normally overccme by the blood clotting mec hanis m tha! causes any gap in a blood
vessel to be plugged. Haemophilia is a rare genecically--determined cOfldition in which the blood
does not clot normally. The result is frequent, =cessive bl=ding.
There are two forms of haemophilia, known as haemophilia A and haemophilia B. 1bey are
due to a failure to produce adequate amounts of particular blood pro!eins essential to the compl=
blood dotting mechanism. Today, haemophilia is effectively treated by the administration d the
dottingfacrorthepatientlacks.
Haemophilia is a sex-linked condition because the genes controlling the production o f the blood
proteins concerned are located on the X chromosome. Haemophilia is caused by a recessive allele.
As a result, haemophilfa is largely a conditiOfl of the male, since in him a single X chrcmoscme
19 Haemophilia i1
carrying the defective alleleex• Y) will result in disease. For a female to have haemophilia. she
5ex-linked condition
must be homozygous for the recessive gene ex• X"), but this condition is usually fatal in utcro. It
that results from the mayre:suh in a natural abortion
inheritanc:eofanallele A female with one X chromosome with the recessive allele and one with the dominant allele
carried on the (X" X ") is a carrier. She has normal blood clotting but for any daughters she has with a normal
Xchromosome male,thereisaS0percentchanceofthembeingcarriers:foranysonsthereisa50percem
Explain why chanceofthembeinghaemophiliac(Figure16.26).
--
haemophilia is more
common in males.
parental(Pl
canierfemale(9) oormalmale(())
offspring(F1)
-
haemophiliac:
~
<<l>
pheno!ype<ratio
371
m 1nheritedchange
Some genes are By the end of this section you should be able to:
transcribed all t he time
toproducecomt itutive a)d istingu ishbetw..enstruct uraland regulatorygenesandbetwe en repreuibleand indudble
proteins; ot hers are only e nzymes
'switchedon'whentheir b) explainge ne ticcontrolofprotein procluctioninaprokaryoteusingthelacoP"ron
protein products are c)expl ain thefunct ionoftranocriptionfactorsingeneexpre .. ionineukaryot es
requ ired d) explain how gibbe rellin activates gene s by causing the breakdown of DELLA protein repressors,
which normally inhibit factors that promote t ranscription
Whena ndwhygene,areexpreHed
ExpreHeda ll thetirne genes,l'5j)Oll5ibteforroutine, rnntinoousmetabolic: functions
(forexa mple,respiratkln),commontoallcells,continuous
th,ooghout life
Ex~Hedataselected stageinaill or ascellsderivedfromstemcells aredevek,p ing intomuscte fibresor
tiHuedeveloprne nt neurooes,forexample
ExpreHed o nly in the matun, cell genes ,l'5j)Oll5ibte for antibody pmduct>OO in a mature plasma cell,
afte,thesehavebeenclonl'd
Ex~ssed o n receipt of a n in ternal or when a partirnlar hommne signal, metabolic: signal, o, nerve
externa l sig na l impulseis,ec:eiYedbythece!l,suc:h as thegenefor ins.ulin
product>OO in flcells in the islets of Lar,gerh~ns
372
16.3 Genecontrol I
How is this regulation brought about?
Regulationofgeneacrioni.spanlyregulatedbythegenesthem.selves.WhUsctheroleofmanygenesi.s
to code for specific enzymes or proceins requira:l by working cells at some stage, others are different -
theirrolei.sroregulateo<hergenes.Consequendy,wecanrecognisetwocategoriesofgene:
• Slnlc turnl gen es code for a structural procein, or an enzyme, or an RNA molecule nor involved in
regulation. They are requira:l by cells to ere-ate or maintain the sirucrure, or en.able the funcrioning,
of the cell or organi'lm. They are transcribed imo proceins.
• Regulatory ge n es are involved in controlling the =pression of one or more genes. They may
code for a proiein or for an RNA molecule.
Genes are also regulated by 'signals" from the environment - typically frcm within the cell. lbese
acr on the enzymes or on the genes that code for them. Consequently, we can recognise two
categories of enzymes:
• Indu cible e nzymes are producn:I under specific conditions. such as the presence of a panicular
substrate. This substance controls the expression of one or more genes, saucrural genes. involved
in the metabolism o f that sub.stance. In the absence of the subsirate, gene action is switched off.
• Reprc,isiblc enzym es are generally produced continuously, but their producrion can be halted.
lbey are formed unless a signal, such as an =cess of producr. turns their producrion off.
373
m 1nheritedchange
rodiog<egnenl
(olexoosaod;mroo,)
I
_Qi:_.
complex(lactors+eoryme)
ootr.epromote<
ThelaN<,rpanofthelig.,reshowsaportioooltheupparDNAmolec:uO,wilhtr.eac1;,aror(0<ange)rpgioo.pmmo1or
(green) fegtOO and codlr,g (hue) rpgion . The DNA is looped so !ml it comes ;mo cootact with l>fferem pans of tr.e
tramcriptooo;niti.nionromplex.Thecomplexcoosiruol"-"'"'"'pmtein,and otherfacttnhence!hemulti?k,
~~~~~tue~"'.!"'r:!~~"::~:~.partoi thecom?k,x. n-e..- ,phefl'<hiWe been <Kawn !Jan,pa,ent
Figure16.l8Coo!Jolsitesandtheinitiationoltran,cfiptioo
374
Summa~ •
How gibberellin activates genes
Gibberellin (GA) regulates and promoces many aspects o f plant growth and development
(page 336). Remind yourself of some of the roie.s of GA now
It is now known that a significarn cause of restraint on many aspects of plarn growth and development
isthepresenceofagroupofnudeus-basedproreinscalledDEILJ.proreins.Thesepro1einsarecoded
for by specific regulatory genes, and they function in the nucleus itself as powerful tran.s,:rip1:ion
factors. How these proreins regulate transcription is poorly understood, but it has been shown that,
when GA levds rise in a plant, DEl1.A proteins are de-activated and the plant grows. For =ample, GA
has this effect as ittriggersexpres.sionofgenescoding forhydrolyticenzymesthatcotwertstarchsiores
toglucoseingenninatingseedssuchasmaizeandbarley(Figurel5.26,pagc339).
The way that GA controls the levds of growth-repressing DEil.A pro!eins is apparently by
labelling them with a molecule known as ubiquitin. With this marker attached, a DEllA protein is
identified and prompdy destroyed by a nucleus-based compl=, which has the role d degrading
unneededordamagedpro1einspresentinthenucleus
• Geneticsisthestudyof inh e ri ta nce.Manycharacteristicsof • Sex is determined in humans by the sex chro m oso mes: XX in
organi:smsarecontrolledbythege neswhich occurasa linear the female and X'( in the male. The X chromosome is longer
series along ch ro moso mes. Diplo id o rga nis ms contain two than the Y and has genes not present on the Y chromowme
mpies of each gene in each of their cells. Alternative forms of In the male, recess.ive alleles on the single X chromosome
agenearecalleda ll e les .lnsexualreproduction, ga m etesare cannotbemaskedastheyoftenarebydominantalleles
formedcontainingonecopyofeac:hgene.Ofupringformed inthefemale.Consequently,anumberofrare rec:essive
fromthezygoteresultingfrom fertili satio n receivetwo cond it io ns are sex-linked and occur more frequently in males
mpiesofeachgene{twoalleles},onefromeachparent Theseindudered-greencolourblindness,haemophiliaand
• Meiosi:si:sthereductivenudei!rdivi:sioninwhich,aftercell Duchennemusculardystrophy.
dMsion,thenudeuspresentineachofthefourdaughtercells • Exceptions to the monohybrid ratio arise from alleles which
formed has half the chromosome number of the parent cell, arecodo min a nt{ratherthantherebeingonedominantand
typicallyoneofeachtype.Meiosisi:saS50Ciatedwithsexuill onerecess.iveallele}.lnaddition,theexistenceofmorethan
reproductionandtheproductionofgametes. two alleles for a gene, known as multi ple a ll e les, complicates
• Alleles may be domina nt or recessive or show todomin a nce inheritance. The human ABO blood group is an example of the
An organism with two identical alleles of a gene i:s ho mozygo us latter.ltisdeterminedbytwoofthreepossiblealleleswhich
forthatgeoe;anorganismwithdifferentallelesofagenei:s givesfourpossiblephenotypes,bloodgroupsA,8,ABandO
heterozygo us. Thegeoeticconstitutionofanorganismisits e Ad ihyb ridcrossconc:ernstheinheritanceoftwopairsof
genotype.Theresultingappearanceoftheorganism,itsvi:sible contrasting characteristics located on different chromosomes
and measurable features, i:s called its phenotype The results confirmthateachalleleofa gene i:sequallylikely
e lnamono hyb rid crosstheinheritanrnofacontrasting tobeinheritedwitheachalleleofanothergene,areflection
chari!Cteristiccontrolledbyasinglegene(suchastalland of the in depend e nt asso rtm e nt of chromosomes that occurs
dwarfheightingardenpeaplants}i:sinvestigated.\/v"hen
parentshomozygousforacontrastingcharacteristicare e Thee nvironm e nta l co nditionsexperiencedbyanorganism
crossed,thefirstgeneration(F,}willbeheterozygouslhe mayalsoinfluenrntheexpressionofalleles.Ageneticallytall
chari!Cteristic they show, rnch as 'tall' in the pea, will establish organismdeprivedofsufficieotnutrieotsandsodevelopsa
thattheallelefor'tall'is d o min a nt overtheallelefor'dwarf', pheootypethatisshortisanobviousexample.Characteristics
whichis recessive.When theF,generationself-fertilise{or influenced by the environment may show a degree of
siblingcros5esaremadeinanimals},arecessivecharacteristic continuous variation.
re-emergesinaminorityoftheoffspring,intherationof3:1 e Mutati o nsmayalsoaffectthephenotype.Whenthenucleotide
(providedmanyoffspringareproduced). Thi:sshowsthatonly sequence in the DNA of a chromosome is changed it may lead
onealleleofageneiscarriedinasinglegamete. toalterationintheaminoacidsequenrnofproteinsformedin
• Since the genotype of an organism showing dominant the cell. Theconsequencemaybeadifferentphenotype
characteristics may be homozygous or heterozygous • Some genes are transcribed all the time, whilst others are only
for the gene concerned, the genotype of this organism switched on\\flentheirproducts are required. Reg ul a t ory
must be determined by a test cross with an organism genescontrolthetimingoftheexpressionofothergenes
thatishomozygousrecessiveforthecharac:teristic.lnthe whoserolesaretocreateandmaintainthestructureor
monohybrid test cross the outcome is a ratio of 1 : 1 and in the functionsofthecellororganism
dihybridtestcrosstheoutcomeisaratioof1:1:1:1
375
m 1nheritedchange
3 a) Explainwhatismeantbyagenemutationduetobase
1ub1titution [4]
b) In the inheritance of sickle cell anaemia, the normal
alleleisrepresentedby Hb andthesic:klecellalleleby
Hb' . Draw and complete a genetic diagram to explain
thegenotypesandphenotypesandtheproportionsof
eachre1ultingfromacrossbel\\!eentwoindividualsof
genotype HbHb' . [8]
Fig.1.1 c) Astudyofalcx:alpopulationinacentralAfricancountry
foundthatinasampleof 12S000people
a)ExplainthemeaningofthetermsaJ/e/eandrece5sive [2] 9449 were homozygous for normal haemoglobin (Hb Hb)
b) Explainwhyfemalesarelesslikelythanmalestohave
RGC. [2] 3020hadsicklecelltrait{HbH b')
c) WithreferencetoFig.6.1,andusingtheiymbols Rfor
thedominanta lleleand r fortherecessiveallele,statethe 31 hadsicklecellanaemia{Hb' Hb'}
genotypesoftheindividual1 1,4,6and7 [4]
(Total: 8] Commentonthedistributionofthesicklecellallele
within this population, and on any environmental factors
(Cambridge International AS" and A Level Biology 9700,
that may influence it [8]
Paper04 Q6June2008)
[Total: 20]
2 In mice there are several alleles of the gene that controls the 4 a) Listfourdifference1betweentheproces1e:Sofmitosisand
intensityofpigmentationofthefur. meiosis,andstatebrieflythesignificanreofeach. (8]
TheallelesarelistedbelowinorderofdominancewithCas b) Where would you expect (1) mitosis and (1i) meio1is to
themo1tdominant occurinafullygro'Nflfloweringplant? [2]
C :full colour c) For a cell with l\\'O chromosomes draw diagrams to show·
i) metaphaseofmitosis
C"'=chinchilla ii ) metaphaselofmeiosi1
iii)propha1ellofmeiosi1
Ch:himalayan
iv) metaphase II of meiosis [4]
d) Oefinetheterm'phenotype'andexplainwithreferenre
C'=platinum
to an example, how the environment may affect
c • :albino phenotype. [4]
(Total : 18]
376
A Level
17.1 Variation
377
m Selection and evolution
~arf:ndtallp• ~•Jon
Theheightsoftheseplaotslallintotwodisc:reteg,oops,iobotholwhichthereisa
.if'°Ua
rftal
oormaldimibutioootvariatioo,bu1withno0Yeriapt,em.,..,,1heg,oops
I
,, :: :'
. ': i '
--
'
' ' '
16 M n ~ ~ ~
(Thesameoumberotdwarlaod~ lplan!<W..ll'meastJ!l'd.)
Hun,.anhe ight·ation
~"
Therffl1,torvaria1iooi11 [30
;~~~~~~n:
dimibution .fo<!hepurpose
i
g
20
oltheg~.!heheightsare
coHectedintoarbio-aryg,oops,
eachofaheigh!rangeo!2cm
height(in2cmbands)
Figure17.1Disc:ootinu0<.1Sandc0<>Jnuoo,variatioo
378
17.lVariation .
The colour of human skin is due to the amount of the pigment called melanin chm is produced
in the skin. Melanin synthesis is genetically comrolled. It seems chm three, four or more separately
inherited genes control melanin production. lbe outcome is a continuous distribution of skin
colourfromverypale(presenceofnoallelescodingformelaninproducrion)toverydarkbrown
(all 'skincolour'allele.scodingformelaninproduction). ln ouriJJustration ofpolygenicinheritance
of human skin colour, we have used only two independent genes. This is bemuse dealing with
say, four genes becomes impossibly unwieldy, on the printed page. Also, the principle can be
demonstrated dearly enough using just two genes (Figure 17.2)
F1 generation AaBb
medium
xb<eedingwi!hirdvidual
ol<ameskinc:OOM"geootype
F2 generation
rang e of AaBB
genotyi-: AAOS
rang aof
phenotypes:
Figure 17 .2 Humanskinc:olou,asacharactefisbc:controltedbytwcindependentgenPS-an;llu,trationof
polygenicinhefitance
379
m Selection and evolution
A further, striking =ample of the inOuence of environment Ofl adult form occurs in the honey bee
(Apis mellifem). In a colony of honey bees there are thr= phenotypes (workers. drones and
queen), but only two genOl:ypes. This was shown in Figure 16.23, on page 366. The drones are
the o:immunity's males and they develop from unfertUised eggs (their genotype is haploid). The
quen, and the workers develop from fertilised eggs and have identical genotypes. The queen, who
is a much larger organism. differs from her workers only by the diet she is fed in the larval stage
(an environmental factor). Her proiein-rich food, prepared for her by the nurse worker bees in the
colony, is not available to the larv-ae that will be workers. This makes a hugely signifirnm difference
intheirphCfX>types.
380
17.lVariation .
between the two darn sets. An example will illustrate the method. However, you should noie that
youarenocexpectedtocalculatevaluesdl.
26,26
J0,31,31,31,32,33 33,34
37,38 35,35,36,36,36,37
41,42
lbe role of the Slatistkal teSI is ro determine whether to accept or reject the null hypothesis. If it
isreje,;:tedhere,wecanhaveconfidencethatthedifferenceintheleafsizesofthetwosamplesis
statistically significant.
Next, check that the data are normally distributed. This is done by arranging the data for sun
leaves and shade leaves as in Table 17.1 (and plotting a histogram, if necessary)
,J - squarerootof
:i Once a value of r has been calculated (the value of I here is 2.10) we determine the degrees of
freedom (df) for the two samples, using the formula·
df = (totalnumberofvaluesinbol:hsamples) - 2
= n. + n,, - 2
lnthis case,df = 11 + 11 = 22.
381
m Selection and evolution
""
'
'
"'
""
""
Figure17.4CriticalvaluesfortheHest
4 A table of Crltka l v a lues for the I-test is given in Figure 17.4. We look down the column of sig-
nificana, levels (p) at the O.OS level until you reach the line corresponding to df = 22. You wUJ
seethathere.p = 2.08
5 Sincethecalculatedvalueof1(2.IO)exceedsthiscriticalvalue(2.08)attheO.OSlevelof
significana,, it indicates that there is a lower than O.OS probability (5%) that the difference
between the two means is solely due to chance. lberefore, we can reject the null hypo1hesis.
and conclude the difference between the two samples is significant.
6 Fortheexperimenter.thesignificanceofthisstatisticisthereisareasonforthedifferena,inthe-
means. which can be funher investigated and fresh hypotheses proposed.
382
17.2Naturalandartificialselection .
Populatiom of organisms By the end of this section you should be able to:
have the potential to
producelargenumbers a) explainth atnatural,e le ctionoccursaspopul ation,havethe capacitytoproduce manyofupring
ofoffspring,yetthe1r thatcompe te forresources;inthe 'strugglefor existe nce'onlythe individual,thatarebest
numbers remain fairly adapte dwrvive tobreedandpusonthe irall e les tothe nextgen eration
constant year a her year. b) explain, with examples, how e nvironme ntal factors can act a, ,tabili,ing. disruptive and
Table17.2Numbersofoffsp,;ogp,oduced
Honeybee (queen)
Poppy
How many of these offspring survive ro breed them<e/..,s?
Consequently. in a stable population parent generations give rise to a single breeding pair
of offspring, on average. All their other offspring may become casualties of the "struggle'. So.
populations do noc show rapidly increasing numbers in most habitats, or at least. nO{ for long.
383
m Selection and evolution
384
17.2Naturalandartificialselection .
parental territory to establish new territories of their own depending on seasonal conditions and the
availability of suitable conditions elsewhere.
In 1950, a biological control agent, the myxcmatosis virus. was introduced . Initially it wiped out
between 95 and 100 per cent of the rabbits in some are-as. However, rabbit populations eventually
recoveredasresistancerothemyxomatosisvirusspre-adamongtherabbits.l.ater,theintroducrion
of other specific viral dise"3ses helped control populations. but again. the rapid appearance of
resistance has left rabbits as a formidable pest in Australia .
'"
3 Whatannotationscould
youaddtothegraphin
Figure17.7
a totherisingsideto
apeakofapredator
b tothedecliningside
of a predator curve
c tointerpretand
explain the results
1hown?
Figure17.7Peltsreceivedli-omtrappmattheHl>dsooil.ayC~ anyOYera1()(}.yearperiod
385
m Selection and evolution
We can conclude from these three comrasting situations that narural populations have the
potential to increase rapidly in numbers provided they have access to the essential resources
they require. However. the size of a population is eventually limited by restraints that we can
call cm·ironmental factors. These factors include space, light and the availability of food. lbe
imeractions of predators or para.sites with a population may be equally influential in decermining
the size of a population. It is this never-ending ccmpctition that results in the majority of organisms
fuiling to survive and reproduce. In effect, the environment appears to be able to suppon a certain
number of organisms and the number of individuals in a species remains more or less constant
over a period o f time. Competition is a major facr of life for the living world.
Stabilising selection
Stabillsing sckction occurs v;hcn environmental conditions are largely unchanging. Stabilising selection
ooc:'l not lead to evolution. It is a med.anism whid, 1mimairu a favourable characteristic and the alleles
rcsp:nsiblefcrthcm,andcliminatcsvariantsanlabnormalitiestrultareusclcssorhannful. Prcrnblymost
populations undergo.stabilising selection. The-c=mple shoovn in Figure 17.8COfllCSfrombinhroc«ds d
human babies born between 1935 and 1946 in a London hospital. It shows there was an qximum birth
weightfcrbabiesandthosewithooth,vcigOO<heavicrOflightcrwcreataselcctivcdisadvantagc.
lheOOhwe+ghtothumao,~lnfluer,a,dlrfenvlro nmentalfadors
(e.g. matemalnutrttloo.s/OOl:Jnghablls.etc.)ar.dlrflnherltance
(about~).
Whenmorebables!hanave<"9"1ieat"""Ylowandvery~blnhw,,jghts.
!h~OOvk>usly affernthegene pool becauselttffidstoelmlnate genes !or
lowandhighblrthwe+ghts
themaln.,aphsl>owsthe
birth wei ghts of Infant•
bornln • Londonhospttal
1935- 19(G {histogram}
386
17.2Naturalandartificialselection .
Directional selection
Dfrce1 io n a l s electio n may result from changing environmental conditions. In these siruations the
matCJrity of an =i.sting form of an organism may be no longer best suited to the environment. Some
cxher, alternative phenotypes may have a seleaive adv:mmge.
An =ample of directional seleaion is the development of resistance to an antibioiic by
bacteria. Cenain bacteria cause disease and patients with bacterial infections are frequently
treated with an antibioiic to help them overcome the infection. Antibioiics are very widely used.
However, in a large population of a species of bacteria, some may carry a gene for resistance
to the antibioiic in use. Sometimes such a gene arises by spontaneous mutation . Sometimes the
gene is acquired in a form of sexual reproduction bel\veen bacteria of different populations.
A 'resistant" bacteriumhasnoselectiveadvantageintheabsenceoftheantibio ticand
must compete for resources with non-resistant bacteria . But when the antibioiic is present
mo.st bacteria of the population are killed off. Resistant bacteria remain and create the future
population, all of which now carry the gene for resistance to the antibioiic. The genome has
be-en changed abruptly.
antibiotic-,esist,ntgeneacquired ~
by mutation
~~
pf<x= isrepeatedfr,another
antibiotk·lreatedhmt
(foHowir,ganothe,infection
andtfeatment withadifferent
antibiotic e.g. methicillin)
catJSingmu'1ipte.a ntibiotic·
resistantbactefiatoevolve
"
""erusear>dmisuseofantibioticscreatethe 0
perfectbreed inggroundfo,resist,ntbacteria 1992 19g,\ 19% 19982000
deve lopme nt o f multiple-res istant
ln thelongtem>. thedrugsindustryfaces
thechallengeof p,oducir,gnewantibiotic, Stap l,y/ococcus aureus,1992- 2001
faste,thanantibiotic-,esi,tantbacteriaevolve (%olinfecteds.am p~receivedinpathologyl..bs)
Figure17.9Mu~pleantibioticresistanceinbacte,ia
387
m Selection and evolution
Disruptive selection
Dis ruplh"c selection occurs when pmt:icular environment conditions favour the extremes d a
phencxypicrangeoverintermediatephenotypes. A!Jaresult,iti.slikdythatthegenepoolwill
become split into two distinct gene pools. New species may be formed. This phenomenon has been
Ulustrnted by colonisation by plants of mine waSle tips (Figure 17.10). These habitats OOen contain
high concentrntions d toxic metals such as copper and lead Mosc plants are unable to grow on them.
However. you can sec from the phomgraph that some hardy graS11es have colonised the contaminated
soil. We can assume that these species have developed resistance to toxic metal ions. At the same time,
their ability to grow and compete on uncontaminmed soils has decreased. Jn the uncontaminated soil
aroundthetipthegra.sscsthathavenoresi.staocetoroxicmetalionsareabletoflouri.sh.
Grasses are wind-pollinated plants, so br=ding between resistant grass plants and non-resistant
grass plants goes on. When their seeds fall to the ground and anempt to germinate, di.sJuptive
selection may occur. Both the non-resistant plants germinating on contaminated soil and the
resistant plants growing on uncontaminated soil fail to survive to reproduce. lbe result is increasing
divergence of the populations. initially into two distinctly different varieties d grass plant . In time,
new species may be formed.
disruptive selection
''
favour,;twoextremeso f the"chosen"characters
attheexpeoseofintermediate/om,s
Figure17.10Miningwastetiphabitat~nddisruptiveselectKln
-
lllit II
I
II
I
Balancing selection
lndirectionalselectionandstabilisingselectionthereisatendencytoreducevariation. Balancing
selection has completely the opposite effect.
Balancing selection i., a process which activdy maintains multiple alleles in the gene pool d a
population. It does this at frequencies well alx,ve that due to gene mutation. An example of thi.s
i., skklc a,ll 1rait. This i., a hereditary condition that causes a proportion of the red blood cells
circulating in the body to be sickle-shaped. You can sec a sampled red blood cells from the
circulation of someone with sickle cell trait in Figure 6.17 (page 125). This illustration also explains
the mutation by which the gene that codes for the amino add sequenced the beta chain of
haemoglobin i., prone to a substitution of the base adenine (A) by thiamine (1) in a codon coding
for the amino add glutamic add. As a consequence, the amino add valine appears at that point
instead. When the genetic code of thi.s mutated gene has been transcribed into messenger RNA and
then translated at ribosomes. molecules d an unusual form of haemoglobin, known as haemoglobin
Hb' results. The molecules with this unusual haemoglobin tend to clump together and form long
fibres that distort the red blood cells into sickle shapes. In thi.s condition the red blood cells transpco,
linle oxygen and they may even block smaller vessels. In people who are heterozygous for thi.s
allele, Hb Hb', less than 50 per cent of their red blood cells are sickle-shaped. Sickle cell trait i., a
non-lethal condition. However, people with it are mildly anaemic and we might =pect them to be
at some disadvantage in life. However, there are areas of the world where this is definitely not so.
388
17.2Naturalandartificialselection .
The world distribution of malarla is shown in Figure 10.7 (page 197). This disease kills a large
number d people each year. Nocice that in regions of Africa and India malaria is endemic. Here,
sickle cell trait confers an advantage. This is because the malarial parasite Plasnwdio,m completes
its lifecycle in red blood cells, but it rnnnoc do so in red blood cells containing Hb'. So a person
who inherits the sickle cell allele from one parent and the nonnal haemoglobin allele from the
other (a heterozygote) is resistant to the malarial parasite. However. people who are homozygous
for the sickle cell allele (Hb' Hb') have sickle cell anaemia and this affects their survival rate.
So we see balancing sel=tion at work here. Tur" back lo Figure 10.9 (page 199) now and
note the corre/alio,1 between the distrib!<tion oft be sickle cell allele and oft be distribution of
malaria in Africa.
lnanyareatherearestrongsel=tionforcesagainscthehomozygoussicklecellcondition:in
areas where malaria is endemic there is also a strong sel=tion force against people homozygous
for normal haemoglobin. However, heterozygoces have a permanent advantage where malaria
exists. Heterozygotes are better adapted than either of the homozyga1es. Because of this
selective advantage, the sickle-cell condition is an example of balanced polyrnorphlsm
stable co-existence of two (or more) distinct types of allele in a species (or population).
Population genetics
Population genetics is the study of genes in populations. Populations are important to our argument
now, because they are where evolution may occur.
A population is a group of individuals of a species, living close together and able to interbreed.
So a population of garden snails might occupy a small part of a garden, say around a compost
heap. A population d thrushes might occupy some gardens and surrounding fields. In other words.
the area occupied by a population depends on the size of the organism and on how mobile it is,
for example, as well as on environmental factors (eg. food supply, predation, etc.). The bou ndaries
of a population may be hard to define. too. Some populations are fully 'open", with individuals
moving in or out from nearby populations. Alternatively, some populations are more or less
'closed", that is, isolated communities, almost completely cut off from neighbours of the same
species.
Figu~17.11The conc:eptof
"population"
389
m Selection and evolution
=, from generatiOfl ro generatiOfl. Remember, 11n 11llele is Ofle of" number of alrem11tive form, of
A ll e le freque ncy:the a gene that c11n occupy" given locus on a chromosome. The frequency with which 11ny panicular
commonness of the alleleoccursinagivenpopul11ti0flwillv11ry
cx:rnrreoc:eofanyparti(ular When a lkk frcq u c ncic~ of a particular popullltiOfl are investigated they may rum out to be
allele in a population sllltic and unchllnging. Alternatively, we may find allele frequencies chllnging. lbey might do so
quirerapidlywithsucceedinggeneratiOfls,forexample.
When the allele frequencies of a gene pool retru1in more or less unchanged, then we know thllt
popullltion is Slatic as reg:,rds its inherited chllracterisiic.s. We can say thll! the populatiOfl is not
evolving.
However. if the allele frequencies of gene pool of a populatiOfl are changing (i.e. the
proportiOfls of p11nicular 11lleles are altered, we say 'disturbed' in some w11y). then we may assume
that evolution is going Oil. For example. some alleles m11y be increasing in frequency because of
an advantage they confer to the individuals carrying them. With possessiOfl of those alleles the
organism is more successful. It m11y produce more offspring, for example. If we can detect chan ge
in a gene pool we may detect evolution happening, possibly even well before " new species is
observed.
Le1thefrequencyolthedomo!lilf>talele(G)bep.andthefrequencyol 11lerecessi,,..alele(g)beq
Tllefrequencyolalielesmustadduptol.sop+q:1
Thi,meansinacros,.aproponion{o)oltllegametescarriestheGalele.andapmportioo(q)ofthegametes
ea<riesthegaHele
Tlleoflspringoleachgeneraliooa<egiYenbythel\mr,ettgrid
So theproger,yarerespective!y:
f)':f,equencyofGGhomozygote
2pq:f,equencyofGghetero,ygote
q'-:f,equencyofgghomo,ygote
Hardy-W•inb• rgformula
!lthelrpquencyotoneallele(G)isp.aodthelreq.encyofthe011le,ajleie(g)isqthen11lefreqo,enciesofthethree
poss;biegenotype<GG.Ggandgga<e,es""'1iwly(Y".2pqaodrf.
lnthiswi1j.HardyandWeinbe,gdeYelopedthelok,w;n9equatiootodescribestablegene"°""
Figure17.12DerivingtheHardy--We,nbergfDfmula
390
17.2Naturalandartificialselection .
The main problem d finding gene frequencies is that is noc possible to distinguish between
homozygous dominants and heterozygoces based on their appearance or phenocype. However,
usingtheaboveequation,itispossibletocalculategenefrequencyfromthenumberof
homozygous recessive individuals in the population. This is ,/. We can find q by finding the square
rool. The result tells us the frequency of the recessive allele, and this can then be substituted into
the initial equmion p + q = 1 and we can find the frequency of the dominant allele.
• Ex a mple l :
In humans. the ability ro taste the chemical phenylthioca,bamide (PTC) is conferred by the
dominant allele T. Bo!h the dominam homozygoces (1T) and the hererozygote.s ( 1"1) are 'tasters'.
lbe non-tasters are the homozygotes (11).
In a sample of a local population of 200 people in Western Europe,130 (65 per cent) were tasters
and70(35percent)werenon-tasrers.
I""'"
Apply this data to the Hardy- Weinberg formula; we know the value of,/ to be 0.35.
Takingthesquarerootthevalueofq = 0.59.
So the frequency of the non-tasting alleles ( t) in this European population was 0.59.
• Exa mple 2:
The absence of the skin pigment. melanin, is a condition called albinism (Figure 16.25, page 370),
a genetically controlled characteristic. An albino has the genocype pp (homozygous recessive),
whereas people with nonnal pigmentation are homozygous (PP) or heterozygous (Pp). In a large
populationonlyonepersonin IOOOOwasalbino.
Thus the Hardy- Weinberg formula has allowed us find the frequencies of alleles P and p in a
population.
391
m Selection and evolution
We have already noted various ways in which natural selection operates to change the
composition of gene pools. A further example will suffice he-re.
Selective predation of members of the population with certain characreristics that are genetically
controlled, will lead to changing frequencies of cenain alleles. For example. selective predation
of snails with a particular shell coloration that makes them visible in (say) a grassland habitat, but
effectivelycamoullagedinawoodlandhabitat(Figurel7.13)
Thebanded,colot.edshejJ.
ofthesna i Cepe,:,nemor;Jis
a,e mmmoo sights in woods.
hedge,;andgr""'1ands.The
shellsmaybebrown.ps1k
ot~low.andpa,;essupto
f;,,,darkbands. lnwoodland
leaf itte,theshel,thata,e
camou/L:,gedaredark!'fand
morebanded thanlhase
camoufla.gedamong,,-asses
Fig..-e 17.13 Setectivepml•booof
~!11~[1
I!
,nails - naturalse!ectioninoperabon
In a very large breeding population there is every likelihood that the sample of genes coming
togetherinzygoteswUJbefullyrepresentativeofthegenepool,andsoofpreviousgeneratioru
too. On the other hand, in a small population the genes selected may not be representative of
thegenepoolasawhole.
In the event of sudden hostile physical conditions, such as extreme cold. devastating flooding
or prolonged drought, a natural population may be reduced to a very few survivors . On the
return of a favourable environment, the numbers of the affected species m ay quickly return to
normal (typically. because of reduced competition for food or other resources). However, the
new population has been built from a very small sample of the original, 'pre-disaster' population,
with numerous 'first cousin" and backcl'06/l matings (causing fewer hererozygotes and more
homozygotes) and with some alleles last altogether.
3 Fo undc refkc 1.
A small group d a large breeding population may beccme isolated, possibly due to a barrier that
arises within part of the territory, or when a small group wander away, or are carried away by
chanc,, - such as by a l5Unami. If these small samples of the original population are unrepresentative
of the original population, a sudden change in gene frequency will have taken place.
392
17.lArtificialselection .
4 0ther 'disrurbingfaaorsatt:
Emigra tio n/ immigra tion . This may introduce new genes into populations, for example
4 What factors may came
Mutatio n . These are random. rare, spontaneous changes in the genes that occur in gonads,
the composit ion of a
leading to the possibility of new charaaerislics in the offspring.
gene pool to change?
Anifici3l selection is selection caused by humans. This process of selective breeding is us1.mlly
a deliberate and planned process. Artificial selection involves identifying the largest, the best or
themOlltusefulof theprogenyfortheinrendedpurposeandusingthemasthenextgeneration
of parents. lben, continuous removal of progeny showing less desired features, generation by
generation. leads to deliberate genetic change. lbe genetic constitution of the population changes
rapidly. Artificialselectionisan on-goingproces.sto obrninhigheryields,superiornutrientconrent
S Charles Darwin and resistance to disease in very many of today's domestic animals and crop plants.
argued that the great Cbarle5 Darwin started breeding pigeons as a result of his interest in variation in organisms
wealth of varieties (Figurel7.14).ln7beOrigi"o/Specieshen01:edthereweremorethanadozenvarietiesofpigeon
we have produced which, had they been presented as wild birds to an expert on birds. would have been recognised
in domestication
as separate species. All these pigeons were descendents of the rock dove, a common wild bird.
1upportstheconceptof
Darwin argued that if so much change can be induced in so few generations, then species
evolution.Outline how
this is so must be able to evolve into other species by the gradual accumulation of minute changes , as
environmental conditions change and select some progeny and not others
B),· artificial selection, the plants and animals used by humans (such as in agriculture, transpon ,
companionship and leisure) have been derived frcm wild organisms. The origins of anificial
selection go back to the earliest developments of agriculture, although at this stage, successful
practice was not based on an understanding of the theory of evolution (Figure 17.15).
393
m Selection and evolution
Wild sheep or mouflon {Ovis musimon) occ ur tod ay Soaysheepofth eouterH e brides Modern selective breedinghas produced
on Sa rdini a a nd Corsica suggesttou swha ttheea rliest s horte r a nimal swith a woollyfleece in
do mesti cat ed s heep looke d like placeof coa rse ha ir a ndwithmusd e of
highe rfatconte nt.M a ny bree ds have lost
th e ir horns
Figure 17 .1 SFrnmwildtodomeshcateds.peciesandtheorigir,so/selectM,breed;ngskH!s
394
17.lArtificialselection .
Cows as rum·nants:
Ce llulose, themost a bund a ntorganiccompound,
isamajorcomponentofthedietofallherbivores.
Ma mm a lsareun a bletoproduce theenzyme
cellulase,butmanybacteriacan.Herbivoresexploit
thisfacilityofmicroorganismsinordertodigestthe
celluloseintheirdiet.Cowshaveafour..:hambered
'stomach'. Th e first compartment. the rumen, is a
la rgefermentationtankcontainingmicroorga nisms
a bletodigestcellulose.Digestionofce llulose be gins
withgrindingbyjawsandteeth.Lateron,inthe
cycleoffermentation,granisalsoregurgitatedto
the mouth from the rume n for furthe r grinding.
Meanwhile, much of the sugar released by cellulase
duringferme ntationintherumenisturnedinto
organicacids.Sugarsa ndorganicacids are the major
sourceofenergyforthecow,andueabsorbed
assoon as theyare f rmed.The m· roorgan·sms
prese nt a lsosynthesisetheirownaminoacids
a ndproteinsfrominorganicnitroge n(- N01 ions,
NH,l.Wh e n.la te r,th econte ntsofth e rume n
pass ontothe tru estoma ch, thecow digeststhe
microorganismsas a n a dditional source ofprotein.
Many cows now produce more than 20000kg of milk per lactatiOfl. This has been achieved
by selecting bulls from high-yield herds. and breeding them with cows that have the best milk
production. lbe use of anifidal in.semiruuion techniques and the maimenanre of stocks of frozen
semen have permitted the use of semen from the most promising of animals to have the widest
impact. However. at the same time there has been a substantial reduction in fenility in milking
herds. Each lactatiOfl starts following the binh of a calf. but first the cow has to be insemin:ited as
the previous lactation comes to an end. This cycle has staned to become problematical in some
of the highest yield cows. Research now indicates there may be a genetic connectiOfl, and the
problemisunderimensiveinvestigatiOfl.
395
m Selection and evolution
Bangladesh
popularion>1610C"OOOO
plommottorindielS = 96%
lcereal, = 82.1%) popularion>62500000
plommonerindielS = 63%
lcemol , 21%)
Table17.3Theworld"stopten
foodaopsandtheesbmated
edibled,ymatter{inmi llionsof
tonnes)theypro...-de
lntraditionalbreedingprojects,themostusefuloffspringforapanicularpurpose,suchas
resistance to specific pest and diseases, are selected a nd used as the next generation of parents .
Offspring regarded as Jess useful are =eluded. This technique has been applied with increasing
sophistication, and is still in common use by researchers.
396
17.lArtificialselection .
....._"'..... There are two classes of wheat varieties . llard w hca ts are higher in
proieins, are grown in areas of lmver rainfall . and are used to produce bread
:.,,,:... that keeps . Soft w h e m varieties are more starchy. are used to make 'French
bread" and pastas, and are grown in more humid conditions.
Productivityinthesecropshasbeenenormouslyincrea...d(Figure 17.18).
This has been deliberately brought about by improvements in cultivation, by
thewidespreaduseof pesciddes,thecarefultargetingofartilicialfertilisers,
1945505560657075 808590952000 andbygeneticnxxlifications.For c xample.theincorporationofmutantalleles
~"
Figure17.18Productivityolwheat
for gibberellin synthesis (the se prevent the action of gibberellin) has led to
new dwarf varieties that put a greater proportion of energy into grain, at the
=penseofstemlength
Plants o riginally grown in the wild would have had some of these characteristics. but not all .
Selective breeding has been undertaken over the years to achieve the quality and consistency
required. Remember, consistent inbreeding (producing generations by self-pollination) leads
eventually to plants that lack vigo ur and fenilicy and have reduced size and yield. This is
referred to as inbreed ing d e pressio n . However, inbreeding does generate plants which
show little variation (a scandardised crop). for they will tend to have the same alleles for
most genes.
397
m Selection and evolution
Wheninbredlinesarecros.sedthe
resuhsarehybridswhichexhibit h}"brid
,igour - the hybrids show greater yields and
more vigorous growth. Thisisbecause,in
them,fewerrecessiveallde-sareexpressed.
However, if cross-fertilisation continues at
random. the re.suits are crops with much
variation betw=n individual plants.
Themostsucces.sfulbreeclingexperintents
havestartedwith"parent"generationswhich
areselectivdyinbredtoddiverhighyiddand
inbredO othervaluablecharacteristicssuchasdi.sease
resistaoce(achievedbya:posuretothepest
,ingle<ro.,,eed ~ •ingl<K:,..., ..... d anduseinfunherbreedingprogramnte-s
IA><Blp,odl>Ced
inonioola1edfield O
(CxD!produced
inonioolot..dlield
only of those pl:int.s which remain pest free)
These parent generations are then involved
indouble-cros.shybridisation(Figure 17.20).
Theresulthasbeenmairefruitthatwhen
planted andcultivatedappropriarely. produces
cropswithspectacularlyincre:asedyidds,
butwithoutthelossdOl:herfavourable
398
17.3Evolution .
Isolating mechanisms can By the end of this section you should be able to:
leadtotheac:rumulation
of different genetic: a) statethegeneraltheoryofevolutionthatorganismshavechangedavertime
information in populations, b) discuss the mole cular evidence that revea ls similarities betw~ do..,ly re lated organisms with
potentially leading to new refere nce tomitochondfialDNAandproteinseque nce data
~cies c) explainhowspeciationmayoccurasaresultofgeographical..,paration(allopatricspeciation),
Over prolonged periods andecologicalandbehavioural,eparation(sympatricspeciation)
of time, some ~ies d) explain th e ,oleofpre·zygoticandp<><t·zygoticisolatingmechanismsintheevolutionofnew
have remained virtually species
unc:hanged,othershave e) explain why organisms become extinct, with refe rence to climate change, competition, habitat
changed significantly and
los.sandkillingbyhumans
many have become extinct
Tab le 17 .4 Charle'.10aiwin'sidea1abouttheoriginofspedes,s.ummarisedinfowstatemenl>(S)andthree
deduc:tions(D)fromthes.estatements
O,gani1msp,oduc:eafa,g,eaternumberofprogenythanevergiverisetomatureindividual,
Thenumbe,of ir,d;vidua!sinspedesrem,fr1S
The re fore, the re mustbe a highmorta lityrate.
Theindividual,;naspedesareootall identical,but1ho\\'Var"1tions;n1he;,d,a,iK!eli1tics
The re fore, some variants will have more success tha n others in the com petition for
survival.Sotheparentsforthenextgenerationwillbese lectedfromthose me mbers
ofthespeciesbetteradaptedtotheconditionsoftheenvironment.
Heredrta,y resemb!ancebetweenparent1andoffspringi1afact
The re fore,s.u bse que ntge ne rationsw illm a inta inandimproveon the de greeof
a daptationofth e irparentsbyg radua lch a nge .
399
m Selection and evolution
Neo-Darwinism
01:lrles Darwin(andne-arlyeveryoneelseinthescientificconununityofhistime)knewnothing
about Mendel's work on genetics. Chromosomes had not been reported, and the existence of
genes, alleles and DNA were unknown.
Instead , biologists generally subscribed to the concept of 'blending inheritance' when
mating occurred (which would reduce the genetic variation available for natuf3l selection).
Neo-Darwinism is an essential restatement of the concepts of evolution by natural
selection in terms of Mendelian and post-Mendelian genetics.
Today, modern genetics has shawn us that blending generally does not occur and that there are
several ways by which genetic variation arises in gamete formation and fertilisation. Neo--Darwinism
is a restatement of the ideas of evolution by natural selection in terms of modem genetics. The
ideas of Neo-Darwinism are sununarised belaw.
7 Putinyourownwords Genetic variation arises by:
the ideas from modern • muta tio n s. including c h rom oisom e muta tio n s and gene mutations
geoeticsthatprovidea • ra n dom assoclll>ent of paternal and maternal chromosomes during meiosis, which occurs in the
basisforthetheoryof process of gamete formation
theoriginofspeciesby • r<.'Combin a tlon of segm e n lS of maternal and paternal h o m o logou s c h rom .-.m es during
natural selection cros.sing over that occurs during meiosis in gamete formation
8 Suggestthesignifican(e • the rando m fusion of m ale a n d fe m a le gam e tes in sexual reproduction (which was
for the theory of understood in Darwin's time).
evolution by natural Then, when genetic variation has arisen in organisms·
selection of the
• it is expressed in their phenotypes
realisationbygeologists
• some phenaypes are better able ro survive and reproduce in a panicular environment, whilst
thattheEarthwasmore
other fail to - a point known as 'differential survival'
than a few thousand
years old? • natural selection operates, determining the survivors and the genes that are perpetuated.
In lime, this process may lead to new varieties and new species.
400
17.3Evolution .
molecules such as DNA, pro1:eins and enzyme systems which is a function of time since particular
organismssharedacommOflancescor.
,~-·-"""'*
tr>e~eate,the -~·
precipitatioo I relatedness to humans. The injected serum causes
theproductiOflofantibodies against the injected
pro1:eins . Then.serumproducedfromthetreated
rnbbit"s blood (it now contains aruibodies against
humanproteins) canbetestedagainstserumfrom
a range of animals. The more closely related the
animal is to humans, the greater the precipitation
observed (Figure 17.21).
TheprecipitationproducedbyreactiOflwith
humanserumistakenas 100%. For each species in
Table 17.6 ,thegre-atertheprecipitation,themore
recentlythespeciessharedacommonancestor
with humans. This technique, called comparative
serology, hasbeenusedbytaxonomiststo establish
phylogenetklinksinanumberofcases.inboth
mammalsandnOfl-vertebrntes.
401
m Selection and evolution
Table17.6Relatednes,imestigatedviatheimmunereact""1
chimpanzee
gorilla
orang-utan
gibbon
'!'idermonkey
We do not know of the common ancestor to these animals and the blood of that ancestor is nOI
available to test anyway. But if the S84 amino acids that make up blood 3.lbumin change at a
constantrate,thenthepercentageimmunological 'disrnnce"betwecnhumansandanyofthese
anim:,ls will be a product of the distances back to the common ancestor plus the difference
'forward" again to any one of the listed animals. Hence the differences between a listed anim:,l and
humans can be halved to gauge the difference between a modem form and the common ancestor.
Since the radiation of the primates is known from geological and fossil evidence, the forward
rate of change since the lemur gives the rate of the molecular clock - namely 35 per cent in 6o
milHon years (my), or 0.6 per cent every million years. This calculation can now be applied to all
thedata(fable 17.6,column5).
402
17.3Evolution .
Figun,17.22TheuseofmitochcmdrialDNA inmeasufir,gevo1utiona,yd;,,e,gerice
Speciation
Present-day plants and animals have arisen by change from pre-existing forms of life. This process
Species: a group of has been called 'descent with modification' and 'organic evolution' , but perhaps 'spt.'Cl:U lon' is
organism1thatare beaerbecauseitemphasisesthatspecieschange. So,whatisaspedes?
reproductively isolated, When Linnaeus devised the binomial system of nomencbture in the 181:h century there was
interbreeding to produce noproblemindefining specics.ltwasbelievedthateachspa:ieswasderivedfromtheoriginal
fertile offspring.Organisms p.tir of animals created by God . Since species had been created in this way they were fixed and
belongingtoa~cieshave unchanging .
morphologic:al(structural}
In faa the fossil record provides evidence that changes do occur in living things - human fossils
s.imilarities,whichareoften
alone illustrate this point. Today, as many different charaaeristics as possible are used in order to
usedtoidentifytowhich
species they belong define and identify a species. lbe three main charaaeristics used are·
• morphology and anatomy (external and internal structure)
• ccllstructurc (whethercellsareeukaryoticorprokaryotic)
• ph}"siology (blood composition , renal funaion) and chcmk~I compoishion (comparisons of
nucleic acid, and proteins. and the similarities in proteins between organisms, for =ample).
403
m Selection and evolution
'Species' or 'variety'?
Since species may change (mostly a slow process), there is a time when the d ifferences between
members of 3 species become great enough to identify separate varieties or s u b.sp,..•dcs .
Eventually these may become new species.
An illustration of this is the development of tolerance to he3vy metal ions in pJams able to
survive a nd even Oourish on the otherwise 1:xire mining waste tips commonly found a t sites where
oresandmineralsh3vebeenmined. Here.he,ivymetalssuch3SZinc,copper,leadandnickelare
often present as ions dissolved in the soil moisture at concentrations th3t generate toxic conditions
for the plants presem on the surrounding unpolluted soils.
We have seen that several heavy metal ions are es.senti3l for normal plant growth, but
only in trace amounts. In mining waste these concentrations of ions are frequently exceeded
and heaps left from 18th- and 19th-century mining acrivities in several countries around the
world remain largely bare of plant cover, even when surrounding, unpolluted soils have dense
vegetation cover. Seeds from these plants regularly fall on mining waste, but plants fail to
However, careful obserVlltions of mining waste at many locations have disclosed the presence
of local populations of plants that h3ve evolved tolerance. One example is the grass A.grostis tcnuis
(Bern grass), populations of which are tolerant of O!herwise toxic concentrations of copper.
Biochemical and physiological mech3nisms have evolved in tolerant species. including:
10 Explain the differences • theabilitytoselectivelyavoiduptakeofhe,i'")'metalions
betweenavarietyand
• the accumuJation of ions that emer in insoluble compounds in cell walls by the fonnation of
a~cies.Findout
stable complexes with cell wall polysaccharides
about an example of
both from an organism • the transport of toxic ions into the vacuoles of cells , the membrnne of which is unable to pump
you are familiar with them out ag:,in, so interaction with cell enzymes is avoided
At u.-hat poi>1/ u~"/1 these copper-n,sis/antforms of Be"/ gross be recognised as a separate species!
The evolution of this form of tolerance has been demonstrated in severnl species of terrestrial
plants and also in species of seaweeds tolerant to the copper in the anti-fouling paints frequently
appliedtothehullsofships
~ - - - IN""tsfrom
unpoluted'iO<I
Figure17.23Coppe,,On
toleranceinpopulabonsof
Bentgras.s
404
17.3Evolution .
405
m Selection and evolution
:::;;:~;=~ . .i~9~;2~itl
~~~n 9 9ti<q9
riverhasfO<fflE'd
406
17.3Evolution .
Manyorgaoism,(e.g. insea,andbi<ck)mayl.Neflownorbeeocarriedoowindnnerrumihe
Galapagos horn the mainland. Mammals are m<l'il unlikely to ha,,.. SI.Jf'lived driftfr,g !hereoo a natural
rafto,,..,thisdis1.10<1',bu!manylargereptilP<canw,viYeloo9period<wW,out food0<wa1e,
Figure17.25TheG.a!apagosls!andsa ndspeciesd"'ergencethere
Another example of behavioural and ecological separntiOfl leading ro spedation has been
demonstrated in the fauna of the Galapagos islands. Today there are 14--15 species of finch,
and tht,>y have all been derived frcm a common ancestor, and have evolved, living in the same
environment. n,.,..._.. birds were noted by Charles Darwin (but they failed to attracr his derniled
attention)whilstonhisvisitstothei.slands. Ata larerdate,detailedstudyofthesebirdswasbegun
by the ornithologist. David lack .
407
m Selection and evolution
lack studied the variation in the beaks of finches - a genetically controlled characteristic. Beak
morphology reflects difference., in feeding habits as members of the local population progressively
focused on different diets, and evolved alternative feeding strategies. lbe evidence they provided
for Darwin"s theory of evolution by natural selection so impressed lack that he coined the name
'Darwin"sfinches". hhasstayedwiththem,misleadingthoughitis.
::,:prourn
~=···
Speciation - a summary
Apart from cases of instant spedation by polyploidy, species do nOl evolve in a rapid way. The
process is usually gradual, taking place over a long period of time. In facr. speciation may occur
over several thousand years. and in all cases requires 'isolation'.
A dcme is the name we give to a smaU, isolated population. lbe individuals of a de,ne are not
exactly alike, but they resemble one another more closely than they resemble members d other
demes. This similarity is to be =pected, partly bemuse the members are closely related genetirally
(similar genotypes), and partly because they experience the same env·-=rnl conditions (which
affect their phenotype).
We have nOled above some examples d the ways demes become isolated Reviewing these. we
see they faU into two groups. depending on the ways isolation is brought about.
• Isolating mechanisms that involve spatial separation are known as a llo p atric s peciation
(meaningliterally'differentcountry')
• Isolating mechanisms involving demes in the same location are known as ~-ympa tric s pcdation
(meaningliterally'samecouncry')
408
17.3Evolution .
Reproductive isolation
By definition, different spedes cannot interbreed and have fenile offspring - gen e flow is
prevented between them. When members of related demes have evolved to this point and have
become fully reproducrively isolated, we recognise them as members of different species. The
barriersthatpreventinterbreedingbetweendoselyrelatedspeciesoccureitherbeforefcrtilisation
can be attempted (pre-zygotic isolation) or after fertilisation has occurred (post-zygotic isolation). In
Table 17.8 the underlying isolating mechanisms are summarised.
Table17.8Repmdtxbveisolatingmechanism
geok;gicaltima'mya
409
m Selection and evolution
The significant forces driving extincriOfls 3re climme cl,3nge. competition between species, whether
iruerspecific or irurn.spedfic, habitat Joss, and by human desa-ucrion
Climate change
Coral reefs are Ofle of the 'theatres of environmental change' thre3tened by global W3rming.
Corals are colonies of small anilllllls embedded in a calcium carbonate shell that they secrete
around themselves . They form their und~ater structures in wann, shallow water where sunlight
penetrates. Microscopic (photosynthetic) 3Jgae live sheltered and protected in the cells of corals.
The relationship is one of mutual 3Jvanrnge (a form of symbiosis called murualism), for the coral
gets up to 90 per cent of its organic nutrients from these organisms. Coral reefs are the 'rainforests
of the oceans' - the most diverse of ecosptems known. Although they cover Jes., than 0.1 per cent
of the surface of the oce:ins, these reefs are home to about 25 per cent of 3Jl marine species.
When under environmental stress (for =ample, high water temperarure), the 3Jgae are expelled
(causing loss of colour). In addition, much of the caibon dioxide that enters the 3tmosphere
dissolvesintheoce:ins . WiththeresultingoceanaddificatiOfl,coralcarulO!absorbtheiOflsthey
need to build or m3intain their calcium carbonate skeletOfls . The coral sc:,ns to die, and the
surrounding marine species, likewise. Mass bleaching events occurttd in the Great Barrier Reef in
1999 3nd 20)2. To,fay, coral reefs are dying all 3round the world . The effects from therm3J stress
arelikelytobeexacerbatedunderfururedimatescenarios.
Figure17.28Coral,.,..fs-thecrisi<situation
Competition
Competition between organisms leading to the =tinction of a species is difficult to esrnbli.sh. We
might argue that any species that has become =tinct and is known only as a fossil may be an
example, but establishing a link between cause and effect may be impassible. Jn the laboratory, we
can demonstrate how competitiOfl between two organisms that have the same diet or nutritional
requirements 3nd live in the same habitat (th3t is, they occupy the same niche) leads to the =clusion
of one (Figure 17.29). This =perimeru is a demonstration of the ccmpetitive =clusion principle.
410
17.3Evolution .
AnexpefimemcamedootbyG.C.Gauseill1934u~ngspeciesoff'ilr"""-'<ll1l,alargepro10,oan
rommoniolreshwater.ltfeedsoo plankto n ,thefoodsotXceusedintheseexpefifflff>ls
P.a<Jrel/a
a smaller,
specle, tulturedsepa rately last-<_jrowing specie,
.
time in clay,
"
specle, cultured tog• ther
411
m Selection and evolution
The history of the evolutiOfl of the genus Homo has many examples of species that became =tinct
(Figure 17.30). Can we know the actual causes of their extinctions? Surely we can only guess.
Figure17.30SomeevoMionary
stages inhom,nidevoluhon
cranialcapad!ylcm'
Habitat loss
Rainforestscoveralmost2percemoftheE3nh"slandsurface.buttheyprovidethehabitatsfor
almo.st50percentofalllivingspecies. lthasbeenpredictedthatifallnon-vertebratesoccu1Ting
in a single cubic metre of tropical rainforest soil were collected for identification. there would be
present at least one completdy previously unknown species. It is the case that tropical rainforests
containthegreatestdiversityoflifeofanydtheworld"sbiomes.
Now. tropical rainforests are being rapidly destroyed. Satellite imaging d the Eanh's surface
providestheevidenceforthis - ifandwherenootherreliablesourcesdinformationareavailable.
The world"s three remaining tropical forests of real size are in Somh America (around the
Amazon Basin), in Wes! Africa (around the Congo Basin) and in the Far East (particularly but not
exclusivelyontheisbndsoflndonesia).
The cu1Tent rated destruction is estimated to be about one hectare (100m x 100m - a little
larger than a football pitch) every second . This means that each year an area larger than the British
Isles Gl milliOfl hectares) is cleared. Whilst extinction is a natural process. this 011Tent rate is on
ascaleequivalenttothatatthetimeoftheextinctionofthedinosaurs(anevent6Smya,atthe
Cretaceous-Tertiary boundary).
412
17.3Evolution .
Destruction by humans
The African ckphant (Lo.xodonla afriama) once roamed masc of the continent of Africa. Jn
1930, it was estimated there were S--10 million African elephants, but by 1979 their numbers
were reduced tol.J million. In 1989, when they were added to the international list of the
most endangered species, there were about 6oo0Cl0 remaining, less than I per cent of their
original number. Although scill relatively widely distributed south of the Sahara, populations are
now fragmented. Many are restricred ro National Parks and R=es. One reason why African
governments take measures to proiect elephants at these venues is the importance of the tourist
trade to their economies. Their National Parks bring in much-needed income. and 'ecorourism' does
not deplete wildlife populations
Today, elephants are threatened by Jos., of their habimts, by conflict with humans and by ivory
poaching.
The de mand for h ·o ry threatens the largest adults with the bigges1 tusks. Old matriarchs (the
oldest adult females who provide the 'social glue' for the herds) are particularly vulnerable. Their
group =isience makes them easier than solitary adult males for the poachers to locate. Other
people continue ro slaughter these magnificent animals too, for the 'bush meat' trade, for example.
Encroachmcnl on human s.elllcn1ents is ana1her problem. The hungry elephant that destroys
crops is often hunted down and killed. Moreover, the increasing pressures of land use, as the wild
spacesaredeforestedorclearedandconvertedrocropproducrion,enhancesthisproblem.lnEast
Africainpanicular,ahighper.:entageofwildlifelivesoutsidereserve~,andsoisnotprotected.
Here, few elephants are now predicted to survive outside high-securiry areas. A similar trend is
seenintherestofAfrica.
The outcome of the5l" threats ro the African elephant is predicted to be =tinction in the wild .
Figure17.31AnAlricanelephanthe«l,observedinaNationalNatureRese,,..,
413
m Selection and evolution
414
Examination style questions I
Examination style questions
1 a)Explainwhatismeantbyartificia/se/ection [4) 2 a) Explain how changes in the nucleotide sequence of DNA
b) In a plant breeding programme, corn, Zea fllilY.,, was bred in mayaffecttheaminoacidsequenceinaprotein. [7]
anattempttoprodureahighyieldofproteininthegrain b) Explainhownat11alselectioomaybringaboutevolution. [8]
TheresultsofthisprogrammeareshowninFig.1.1 [Total : 15]
(CarrixidgetitemalimalASiJndALeve/Bidogy9700,
Paper 04 Q9 Jwe 2009)
41S
A Level
18 and
Biodiversity, classification
conservation
ThebiodiversityoftheEa rth is threatened by huma n activitiesand stude nts to obse,ve different species intheir loca lity a nd assessspec ies
climate cha nge. dassificationsyst e msatte mpt to put orde ron dist ribu tion and abundance. Conserving biodiversityi sa diffic ult task
the cha os of a ll the o rga nis ms that ex ist on Earth. Fi e ld work is a n but is achieved by individua ls, local g roups, national and inte rnational
im po rta ntpart of abiologica l educationtoapprec iate this diversi ty orga nisations. St udentsshould a ppreciate th e th reatsto biodiversity
a ndfindouthowtoa na lyse it.Thereareopportunities inthistopic for a nd considerthestepstakenin conservation,both locallyand globally:
«? 18.1 Biodiversity
Biodiversity is much By the end of this section you shou ld be able to:
morethanalistof
allthespecies ina a) de fine the termsspecie s, ecosyste mandnkhe
particular area b) explainthatbiodiversityisconside re datthree differe ntleve l.:
• variationin ecosystem,orhabitats
• the numberof,peciesandthe irre lative a bundance
• ge ne ticvariationwithin e a,chspecies
c) explainthe importance ofrandom.ampl inginde terminingthe biodive riityofanarea
di un ,uita bl e method., such as frame quadrats, line transects, belt tran..,cts and mark·re le ase·
recapture,toassessthe distributionandabundanc:eoforganismsinalocalarea
e ) useSpearman'srankcorrelationandPearion'slinearcorre lationtoanalyse the relation,hips
betw.... nth e distributionandabundance ofspeciesandabioticor bioticfactori
f) use Simpson'slndexofDiYenity(O)tocalculate the biodiversityofahabitat,usingthe formula
Spec ies:Agroupof
organisms of common
ancestrythatdosely
Species, ecosystem and niche - core concepts
resemble each other
structurally and
in environmental study
biochemirally,andv.t.ic:h
are members of natural
Species
populationsthatareilclually It was the Swedish botanist Karl Linnaeus (1707- 78) who devised the binomial system of
or potentially capable of nomencbrure in which every organism has a double name consisting of a Ultinised generic name
breeding with each other (genus) and a specific adjective (species). There was no problem in Linnaeus· day in defining
to produce fertile offspring, species because it was believed that each species was derived from the original pair of animals
andwhic:hdonotinterbreed created by God. Since species had been created in this way they were fixed and unchanging.
with members of other In facr the fossil record provides evidence that changes do occur in living things . 'Humanoid"
and human fossils alone illustrate this point , as we shall see. Taxonomists now use as many
416
18.IBiodivml'Y I
differentcharactcristicsaspossibleinDfdertodefineandidentifyaspecies.Thethrecmain
~m"dd=~
1 Name one ecosystem
inyourlocalitythatyou
might study and then
~;;tnmpl
listthreeabioticfactors
affecting living things in
that ecosystem
~ oopondbottom
nic heofdetfitus-feed;r,g
fish of pond mud
Nich e:thefunctional
roleorplaceofaspecies
==:=;:inir,g examples of ha bitat s
417
m Bicxliversity. classific.ation and con~Mtion
lbtfish. The shag builds its nest on much narrower cliff ledges. It also feeds further ouc to se-a and
captures fish and eels from the upper layers of the water. Since these birds feed differently and
havedifferembehaviourpanerns.althoughtheyoccurincloseproximity,theyavoidcompetition.
Compc tiiio n refers to the interaction between two organisms siriving for the same resource. For
competing organisms, the more their niches overlap, the more that both will strive to secure a finite
resource. Hence many potential competitors have t,>volved differem niches.
surfac:e-swimmingp<ey {""'°""
herring
lla1fi1h
bonom-leedingp<ey
{ .,.,..,
shri~ .
Figure18.2Diffe,entnkhesofthe1h.tgandthernrmorant
Habitat
The habitat is the locality in which an organism occurs. It is where the organism is normally found.
Habitat:theparticular ltiseasytogivethehabitatdsomeorganisms,forinsrnncetheredsquirrelisrestrictedtolarge
locationandtypeoflocal blocks of coniferous forest in northern Asi3 and Europe. The seaweed bladder wrack is found only
environmentcx:cupiedby inthemidzoneofthetidalseashore.Ontheotherhand,thedandelionoccurswidely - onarange
a population or organism, ofsoilsbutalsoincrevicesofwallsandbetweenpavingstones.
characterisedbyit1 phy~cal If the area of the habitat is exiremely small we call it a m lcroha bil a t. The insects that inhabit
featuresorby itsdominant
the crevices in the bark of a tree are in their own mkrohabirnt. Conditions in a microhabitat are
produc:er1{1ucha1rod:y
shoreorsugarcanef1eld}. likely to be very different frcm conditions in the surrounding habitat.
Figure18.3Thehabrtatsofbladderwrac:karidthedandelion
418
18.IBiodivml'Y I
Population
Population:allofthe
By population we mean all the living things of the same species in a habitat at any one time
organisms of one particular
~cieswithinaspecified lbe members d a population have the chance of interbreeding, assuming the species coocerned
areaataparticulartime, reproduces sexually. All the genes and their differem alleles presem in an interbreeding population
!.haringthesamegenepool is called the gene pool. The boundaries of populations of aquatic organisms occurring in a small
andmoreorlessisolated pond are clearly limited by the boundary of the pond. The populations of the garden snail may
from other populations of befoundincomrastinghabitats. SomesnailsoccurinextensiveandUJ-definedareas. around a
the same species. compost heap , for e xample, whilst others occur in small reSlricted areas . Meanwhile , a songbird
pred:itorofthesnail.thethrush,might occupythewholegardenanditssurroundingfields
and hedgerows. In a1her words . the area occupied by a population depends on the size of the
organism and on how mobile it is, for example, as well as on envirorunental factors, such as food
supply and predation.
So the boundaries of a population may be hard to define. Some populations are fully 'open'. with
individuals moving in or out from nearby populations. Alternatively, some populations are more or less
"closed'. that is, isolated o:mmunities almost o:mpletely rut df from neighbours d the same species.
Figure 17. 11, on page 389. describes the coocept of 'population". I.ooh back at it now
Community
Commun ity: all of the A community consists of all the living things in a habitat, the total of all the populations , in fact.
popul ations of all of the So, for example, the community of a well stocked pond would include the populations of rooted.
different species with in a Ooating and submerged plants, the populations d bottom-living animals, the populations of fish
specified area at a partirular and non-vertebrates d the open water and the populations of surface-living organisms - typically a
time verylargenumberoforganisms
Within any community there exists complex interactions between individuals - necessarily,
organisms affect each Olher. This is because the resources required h)-· plants and animals are in
limited supply. Competitive interactions occur between members d the same species (lntraspcdfic
compctitlon) and between members of different species (inters pcdfic competition) . We will
re-visit aspects d these interactions when we discuss food drnins and food wcboi. energy transfer
between trophic levels and the cycling d nutrients within the environment
2 Usethetermsbelowto
describe one or more
of thefeaturesofa Introducing biodiversity
freshwater lake listed There are vast numbers d living things in the world. The word 'blodh-ers lty ', which refers to this,
(a Hg)
is a contraction of 'biological diversity' . It is a tern, we may use in three different ways.
population ,
ec:osystem , habitat,
ab ioticfa ctor, 1 Biodivenity - the total numben of species known,
commun ity. biomass and their relative abundance
a Thewholel ake Up to now, about 1.7 million species have been described and named. However. until very
b Allthefrogsof the recently there has been no anempt to produce an international 'library of living things" where new
eke discoveries are automatically checked (see below). Consequently. some known organisms may
c Theflowofwater havebeen'discovered' morethanonce.
through the lake Meanwhile, previously unknown species are being discovered all the time. In the UK alone.
d Alltheplantsand several hundred new species have been described in the past decade. We might have expected
animals present all the wildlife in these islands to be known, since Britain was one of the countries to pioneer
e Themudofthela ke the systematic study of plants and animals. Apparently, this is nOI the case; previously unknown
f Thetemperature organismsare stillfrequentlyfou ndhere.
variations in the lake Worldwide, the number d unknown species is estimated at between 3 and 5 million al the very
g Thetotalmassof least, and possibly as high as 100 million. So scientists are not certain just how many different
vegetationgrl'.M'ing types of organisms exist. Figure 18.4 is a representation d the proportiOfls of known and unknown
in the lake speciesestimatedtoexlstinmany ofthemajordhisionsoflivingthings.
419
m Bicxliversity. classific.ation and con~Mtion
protoctista insl'cts(adass
of the arthropod,)
Ofthe1.7millioodesaibedspecies,
mo«'thanS0%areir,seru;,aodthe
t,q,e,J)OOts,mostlyllowefingplarm,
arethenex!!arge,t<7"'4'. By
comrast,only4000specie,ol
m>mmalsa,ek""""',aoout0.25%
dallkoownspecies
,,...,tel,,ates
- fish.amphibians, , keletooandjointed
<"!'liles,birdsa nd limbs)
mammals
Figure18.4Therelatrvenumbe,ofanimalandplantgroups
420
18.IBiodivml'Y I
Investigating aspects of biodiversity
In the siudy of a habitat it may be important to collect accurate information on the size of
populations present. A total count of all the members of a population is called a cen s u s. However.
such an approach is usually totally impractical because of the sii.e of the habitat. the movement of
animals, and the huge numbers of organisms usually involved.
For practical reasons it is a random sample that is selected for this. involving either a pan
of the area of the habitat, or a restricted number d the whole population. By means of random
sampling. every individual of the population has an equal chance of being selected, and so a
representativesampleisas.sured(Figure 18.5).
2C0-0rdonatest0<plac:in9quitdraisa<e
obtainedassequern:esolrnodomnumbers,
us.ingcompu1e,soflware,oracakulat0<,
or published table,
'=•
,~"
'~
00
;o
"
"
0 102030405060708090!00
00 r-t---t-i-t-++-1---t-.t-1
i ~ I'-·+-l-++-t-+-+-+-l---0
3 Wi!hineac:hquadrat,tr>eindMduai,pecM'S
'~H'+±..t-+-H-++-1
{'f . ~ ~. ~.
areidemified,and!hen tnl'density,
lrequeocy,ct1Yerorabondanc:eoleac:h
speciesisestimatedfigure18.6
4 Oensi~lrequency;<DY!'f,orab,mdaoce
estimates aft'then quan~l>edbymeasu,ing
1ne1otalaft'aol1nl'hab<tat(!hea,ea
ocrnpi,edby!hepopulat Klfl)in,quaremetres
Themeaodensity,ffequenc)l;rn""'"'
ab<indaocecanbecakulated,usir,gthe
equation
p<l!l<llabon size,, mean dens:;.!"~/·~:-~~:;~ x total a,ea
Figure18.SRandomkxat,ngolquacfra1'
-
421
m Bicxliversity. classific.ation and con~Mtion
density=meanoombersofiodividualsofeilChspeciesperunitarea(time-con,uming
and maybe tla<dtoa,sess separate individuals)
f requ e n cy=numberofqu,>dratsiowhkhaspeciesor:ru".exp,e,;"-'<las%(rapidand
usefulforoomparinglW<lhabitats)
cover =the % ofg round c0Yeredbya,"°"',es(usefulwhereitisootposs.bletoidentify
separateindiY,duals)
a bund a n ce= subjective assessmentol species po,sem. using t~ DNOR scae: D = dominam,
A=abundan~ f =f,equeo~ O =occasional. R=ra,e(sameobserver must make abundance
judgement,;,....+,ict,maybeusefula,;comp,Yisonsoftwoormo<ehabitats.rather!han
objective scores)
sC<speciesp<es.ent.Howmooydifferemspeciesare
coonted io thequadratofsidesl 0.20.30.40.50. '. 1'.,,, •, .:. '-!•· •,, .. ~
~
~-.:...~:..:dn ~~i,.1.':"nc%"=:~s:~ : ' i j rrt • >
, ";. i i 1 I> ..) V ..
0<ve,yfewhmherspeciesp<eseot
How many quadrats? When there is oo fur1her
inc:reilSeint~numberofspeciesloond,sufficief\1
:· j .j. j· i~ : ·. ·...
quadrarsh..,ebeenanafysedin thathabitat
··· ~~\
1
.° / ·. ~ :t•:.':/
. ;.., '. jj e'1't... • ~ t,
••••• :- • • ,v j . .. . . ....:.
!
u"-'<l.ooadditiooal
40 spec,esa,elound
(inthishabitat)
30
0,0 ._,
i 10
0
10cm
quadrat
c o 10 20 30 40 50 ~,m
numbe,olquadratsanafysed quadrat
422
18.IBiodivml'Y I
Estimating species
distribution by means
of a transect
Whereas some conununities are
relativelyunifonnoveragiven
area and are suitable for random
analysis(suchasthegrns.sland
habitat shown in Figure 18.7),
cc:her:sshowatreodinvariation
inap:uticulardirection.
Exarnplesincludesea.sha-e,
pondorlakernargin,salunar.sh
orevenanareawherethereisa
changefromdrysoiltowedand.
lheappropriatetechniqueto
studysuchatrendofvariatlonis
the transect.
Transectsareameansof
samplingbiO(ic(andabiO(ic)
damatrightanglestothe
impact of unidirectional
Figun,18.7Anareaofcl,alkgrass!,mdwithafichflora physical forces.Although there
may only be time to study one
transect in detail (and this may be sufficient as a demonstration of the zonation of communities), a
single cransect may nO( provide an adequate sample or give an indication of differences from place
to place. Transects should therefore preferably be replicated several times.
Where transects are carried out across a habitat where the land changes in height and where
level is an impon:arn factor (such as a seashore, sahmarsh or JXXld margin), then the changes in
level along the transect line can be measured and recorded as a p rofile tra n sect. The swveying
for this requires the use o f survey poles and a field kvel device (Figure 18.8)
The cooununity presern along a transect can be analysed from a straight line such as a
measuring tape, laid down across an apparently representative pan of the habitat. The positions of
evctyorg:,nismpresentthattouchesthelinearerecordedeitherallthe wayalongtheline orelse
at regular intervals. The result is a line tra n sect.
A belt tran sect is a broad transect. usually half a metre wide. To produce it, a tape measure or
rope is laid as for a line transect, but this time the organisms in a series of quadrats of half-metre
widtharesampledat (say) metreintervals. lfthecooununitychangeslittlealongthetransect.then
quadrats can be placed les.s frequently. Along with data on the biota. data on a bio tic variables can
be measured and recorded along the transect. For example, along a terrestrial transect the soil pH
might be measured.
The results of a belt transect study of a seashore community are shown in Figure 18.8. The
seashore (known as the littoral zone) is a pan of the extreme margins of continents and marine
islands periodically submerged below sea water, and so affected by tides. Tides are the periodic
rise and fall of the sea level due to the attractions (gravitational pull) d Moon and Sun. lhe shore
is an area rich in living things. and almost all are of marine origin.
The higher organisms oc01r on the shore, the longer the daily =posure to the air the organisms
endure. Exposure brings the threat of desiccation and wider extremes of temperature than those
experienced during submersion . 'Exposure' is an abiO(ic faaor that influences distribution of
organisms on the seashore.
423
m Bi .id r,J~ d.mifiation n:I toor.tMtion
plllnts
blacklichen{\.em.lc.uiam.J<1ra)
channea.d,.,ack
(Pet;etjj~Q)
spiralwrlld(Fucu,,pirali,)
--·
~nod:»um)
blackwrlld(hcusve,ku,b"'')
1nlmll l•
n«ltewinl:le(Lil'roma neriroides
i i
roughwinkle(Lll'rorinarudi,)
i i
ed~lewinkle
(Urtorir,a/itro,...aJ - ; - ! •
......
smoolh\ml;le
(Utrorir,aobtuuta)
/Mxellalapilt,s)
:::"'&lgu)
"'""'
(SemiN.litnusba~)
conwnonli'npel(Pa!el.:i'oUgata)
~,
a ahighwaterlocation.
b alowwaterlocation
oftherodr.yshore
Fj,g1.re 18.8 ~•ndb.ittm-.-:tllflilly;isofllrodyshorecommuruty
414
18.IBiodivml'Y I
Mark, release and recapture (MRR) technique (the Lincoln Index)
We have already nOl:ed that an important .step in decermining the biodiversity of an area is the
collection of accurate information on the size of populations preseru in a habitat.
A torn] count of all the members of a population is called a census, and would give us the most
accurate data. However, the taking of a census is normally imprnctical. This may be because of the
size of the habirnt, or because of the quick movements of many animals, or because of species that
areonlyactiveatduskorafterdark.
The cap1ure. m a rk. release a n d r eca pm n, (]\!RR) technique, also known as the Lincoln lnd=,
is a practical method of estimating population size of mobile animals (N), such as small mammals,
woodlice or insects that can be caprured and marked with a ring, tag, or dab of coloured paint or
Of the firs! sample caught for marking (n1), it is es.sential to use a method of marking totally
resistaru to removal by moisture or by deliberate actiOfls of the animals. It must be a method that
does no hann to the caprnred animals, too. For =ample. these must nOI be more visible and
therefore more vulnerable ro predation than normal. It is also essential to trap relatively large
samplesiftheresultsarerobesignificant.
The method then requires that marked individuals, Ofl their release. are free to distribute
themselves randomly in the whole population. After randomisation has occurred. a second sample
is caught (n,), some of which wUJ be from the first sample (niJ, recognised by their marking. The
sizeofthepopulatiOflisestimatedbythefonnula:
N= "•n:"'
MRR may be demonstrated on populations of woodlice discovered shdtering under stones/flower
pots. etc. in an area of a garden or woodland. Alternatively. on populations of night-flying mOlh.s
via light traps, or on populations of small mammals trapped in Longwonh small mammal traps
providedsuffidenttrapsareavailable,for=ample
Prior to experimenting on natural populations, the technique can be learnt and understood
by appHcatiOfl ro non-living models. These may include beans dispersed in sawdust, or by using
a 'population" of dried peas in a hessian bag. Here, samples are taken by hand, the sampler
blindfolded,ineffect. lbesetrialsenabletheaccuracyoftheMRRmethodtobetestedbecause
the actual 'population' size (beans or dried peas, etc.) is easily known. 1t will quickly be evident
that the size of samples captured determine the accuracy of the technique; small samples may g ive
wildly inaccurace estimates. The steps to the MRR technique are outlined in Figure 18.10
4 U1ing MRR, relatively large samples mu:st be caught for significant re§Ults. Cakulate the
estimated 1ize of the population in Figure 18.10 if the second sample had included llm
markedwoodlice,notone
5 As a new habitat was colonised, the size of a population of the common rough woodlouse
Porce/lio Kaber was investigated by mark, release, recapture, with the following results
,.,
C.ptured,markedandreleased(n,l
C.pturedintt.esl!'Condsamph,g(n,l
Markedindividualsrecapturedtn,)
425
m Bicxliversity. classific.ation and con~Mtion
ln"marlc.release.rPCapture" !l>e
sizeoftl>e,,,.imalpopulat>OO
isestimatedbythelonoola
N: thepopulalioo
beingeslimated
'
. n1 : numberofma<ked
individllalsrecapwrl'd
L .
:.::.,
Figure18.9Arethetwo
variablescom,latedl
.
24nDY15 laterasecond
,andoms.,mpleiscaught
intl>es.,meK>Gaiity
Figure18.10£stimatir,ganimalpopulationsusing"mark,,eleaseandrecaptu,e"
426
18.IBiodivml'Y I
Speannan's rankcorrelation Peuson·s~neucorrelation
Forordinal(non-ooonal)data - values0<observationsthatcan Fmnormaldata - valuPS mobservabomthatcanbeassigneda
beran~ed. counted and ordered number (as a label)butdatacanoot be ran~ed
Examples Howdoesthepopu!ationsizeofgrasshowersva,yinll'latiooto Kowdoestfieheight offathm,e\atetoheightofthei,sonsor
wheope<tic&we,e!astapplied - an,fewe,.-.wrnf"'-'"'<lwhere daughtm - dotalle,fathershavetallerchildren?
pPStiode~icationisrecent?
Kowdoesspeciesdive"ityre\atetolatitude - doesspedes
Howdonumbers ofaquabcstoo,..ftyn)fflllhsrelatetothe divmitydec,easewithdistanc:efromtheEquator?
hardoessofthewater - a.remoreoymphsp""'-"11inhardwatefl
Kowdoestfiesize oflanduabs,elatetotfiedemityoftheir
Howdoesthedepthofa s!Team,..\atetotheproximityofthelM.'f bo1TOw - aretheburrowsollargec,absmo<edispersed?
banl< - doesdepthofwaterir,aeasewrthdistancefmmtheedge7
427
m Bicxliversity. classific.ation and con~Mtion
Sanddepositedbytheseaandbiownbythev.ind,
b<Jildsintosma l heap,arO<K>dpjooee,XiM>phytk
plar,ts,suchasmarrarnandrouchgras,,atcoam
whefethep<eVai lir,g v.ind;,111>-sho«>.Thetuftsol
H,ave:1growir,gthroo~thesaridoccele,a1eciepositioo,
andgradualydriftin9<ar>dsga1hefadensec0Yerof
vegetatioo. l1Xiedsariddu"'-"aft'looned
polntframo quodr., ln UM
Theframe;,randomly?'Kl'dalargenumberof
times,the10pimloWl'fl'llintumornovegetation
ar>dthespecies(orbaregroor.d)reronled
Studyof a sanddune su«essionfrom e mbryostate tofin ddun e
'=""9'""
,pear-H,avedaoche
pricklysal!Wort
bitingstooecrop
buttercup
stag;hom?"f\tain
v.ildthyme
428
18.lClassif:ation I
4' 18.2 Classification
Organimis studied locally By the end of t his section you should be able to:
may be used to 1how how
hierarchical classific.ation a) describe th e classification of species into the taxonomic hie rarchy of domain, kingdom, phylum,
systems are organised class, order, family, g=u• and species
b) outlinethecharacteristkfeatureofthethreedomainsArchaea, Bacte ria and Eukarya
c) outline the characteristic f.,..tu,.,. of the kingdoms Protoctista, Fungi, Plantae and Animal ia
d) explain why viruses are not include d in the three domain classification and outline hew they are
classified, limited to type of nucleic add (RNA or DNA) and whe thf'r th""e are single stranded or
Europea n mag pi e {Pica pica) As ia n magpie {P/,. tysmuru, /eucoptenJJ} Sri Lanka n mag pie (l.b'oo'ua ornat e)
So, each organism is given a scientific name consisiing of two words in latin. lhe firs1 (a noun)
designate, the b"'""", the second (an adjective) the species. lhe generic name comes fir.st, and begins
with a capital letter, followed by the specific name. By convention, this name is wrinen in italics (or
is underlined). As shown in Figure 18.13, closelyrdmedorganisms have the same generic name; only
their species names differ. You wm see that when organisms are frequently referred ro the full name is
giveninitially,butthereaftcrthegenericnameisshonenedrothefirs1(capital)lettcr.Thus,incontinuing
references ro humans in an an:icle or scientific paper. Homo sapiem would become H. sapiens.
429
.-._
m Bicxliversity. classific.ation and con~Mtion
Homol>ab ili, {ha ndy Homo sapiens {mode rn huma n} Pa nthera /eo Oionl Panthera tigris {tige r)
ma n -ex tinct)
8 Scientificnamesof
organi,m,areoften
difficulttop-onounce
and to remember. bµain
why they are used.
'w I
Figure 18.13 Nam;r,g organem, u,;ng the binomial <)'Stem - ge<>eric name (noun)-+- s.pec;fK name (adjective)-+- common name
kingdo m !helarge,tandmostinclu,.;.,,.
d u, agmupingoforc!ers
•""'" l"""m
orde r :~~!';:"""rty
J
...;.~:::::t "':"'="'
genusagroopof5'mila,and
'"'"'"T""'"
speciesagroopoforganism<
capable olin!erbrl'l'dngto~e
fertileolfsp<ing
Figure18.14Thetax.ausedintaxonomy (a,memonic:to,emember!hehterar<hyolt=
andappliedtotwodifferentkingdoms Ki1>g Pe!ercalledOut forGenui,,...Sder\tiru)
430
18.lClassili:ation I
Domains and kingdoms
At one time the living world seemed to divide naturally into two kingdoms COflsisting of the plan ts
(with a m o trophk nutrition) and the a nimals (with h ctcrotrophk nutrition). lbese two
kingdoms grew from the original disciplines of biology. namely botany, the srudy of plants, and
zoology, the study of animals. Fungi and mkroorg:misms were conveniently 'added' to bornny!
Initially there was only one problem; fungi possessed the typically 'animal ' heterotrophk nutrition
butwere'plant-Hke' instrucrure.
Prokaryotn thepmkaryotekir,gdom,thebacteliaandcyaoobactelia{pl,otosynthetkbac:te,ia),
predom;nate!yunic:ellularorgan;.;ms
theprotoctistank ir,gdom,(euka,yotes),piedominatelyunkellula<,andseenas,esemblir,g
theancestorsofthefungi, plants and animals
Fungi the fur,gal kingdom, (eukaryotes), predominately multkellula, organisms, non-motile, and
withheterotrophknutrit>Ofl
the plant kir,gdom, (eukal)'{lles), multicellular organ;.;ms, non-motile, with autotrophic:
Notethatvi,usesarenotdassmedaslivingorgan;.;ms - seebek>w.
Halophytes - 'salt-loving' bacteri.a commoo in salt lakes and "tiere sea water becomes trapped and
concentratedbyevaporation\\tieresalthasc,ystallised
Alkalinophiles-'alkali-loving'bacteria su!VM'ataboYepH10,conditionstypicalofsodalakes
Acidophiles-'acid-loving'bacteria bacteliaofextremelyacidiccooditions
Them,ophiles-'heat-loving'bacteria oc:cu,inhot-springsatabout70<>C.Someareadaptedastosu,vive
attemperaturesof100-115°C{hyperthermoph ilicpmkaryotes)
Cryoph~• - bacteria of sub.zero common at temperatures of-10<>(, as in the ke of the poles
temperatures ,"'1eresaltdepressesthel,eezingp0<ntof"·ate,
Sincethesenameswereliistused,theseorganismshaw,been/oundinabroaderrangeofhabitats.For
example,someoc:curintheoc:eans, andsomeinfossil-fueldepositsdeepunderground.Somespeciesoc:cur
onlyinanae,obic:environmentssuchasthegutso/tem,itesandofcattle,andatthebottomolpoodsamoog
the rntting plant rema ins. Here they break down organk matte, and ,elease metha<>e - with ve,y important
en ·ronmental consequences
431
m Bicxliversity. classific.ation and con~Mtion
•n,.,st,oo.,1branc:M'Sleitdtohypenhe<rnophilic:spedl'Sv,hichsugges1'thatthe
u,.,.,.,....,.lanc'-"'lorofallli,,;ngthingswasahypenhe,mophile(possibly'as"-'fnbled'
underconditioo,atdeepor:eilllvm!swherevokallkgasesaredi,c:ha,gedintowater
athigh!emperaturear>dp,es.sure)
An:haea w ere discova r«I among prokaryotes of u trem• and in aa:assibl • habitats.
Subsequa ntly, otha r members of th• An:haea ware found more widaly- in th•
gutofha rbivoresandatthebottomoflakasandmountainbogs,foruample.
Figure18.15RibosomalRNAandthedisw,eryofarchaea
432
18.lClassili:ation I
Al l organisms are now dassifoid into three domains
As a result, we now recognise three major forms of life, called domains. lbe organism., of each
domain share a dislincrive, unique pattern of ribosomal RNA, and there are other differences that
establish their evolutionary relationship., (Table 18.4). These domains are:
This major advance, the recognitko that the Archaea ,vere a separate line of evolutionary descent
from bacreria, was made by Carl Woese in 1977. You can read about him if you put his name into
a search engine.
Eukaryota
DNAofchromosome(s) c;,rnlargenome circ:ularge
1boundprote;n(histooe)p<e<ent present present
Lipidsofcellmembranebilayer branescootafr,lipid,thatdifferfromthoseofeubacteriaandeuka,yotes{Figure18.16)
Pho•pholipid,ofarcha.alm..,,,branH
9 In Figure 18.14t\m
animal species are
classified from kingdom hydrocarboo ta~,
to species level.Suggest
how this flow diagram
~ (- 0
needstobemodifiedto
show its classification ~ ,-
lrom'domain'level ethe,hrl<a,~/ Phospholipid,of<1ubact<1ria
and<1ukuyot.m<1mbranH
~droca,bootails
0-<jyceml
- ~~
/'v'v~/vvv,,r
esterlirl<age/
phosphate group
Figure18.16Theli!)idstructureolcellmembranesintheth=domains
433
m Bicxliversity. classific.ation and con~Mtion
~=:::-~
,...~-"'··~-...
""''
~-~
- -
li<,! 01--S<flsitN<
nucleus
:::..
·-- ~-~=. me<ro<..,.
0
t>m>sD"9<
-N
Figure18.17 TheProtoctista
434
18.lClassif:ation I
The Kingdom Fungi
Includes the moulds, yeasts. mildews, mushrooms and bracket fungi.
Thech3ractcristksoftheFungi·
• l>lulticdlubr organisms (excepi: the unicellubr yeasts), not differentiated into tissues and non-mo(Ue.
• Cells are eukaryotk. with a distinet nucleus and membrane-bound org3nelles.
• Nucelus contains linear chromosomes of DNA with histone protein anached.
e largeribosomes(80S)present.
• Nutritionishereruophk.
• Bcxlyis tru1de of branching, rube--Jikehyphae, oftendividied bycros.s-waUs into multinudeatesection.s.
e Reproducebyspores,producedbyasexual3ndsextl3lprocesses.
Yeastfeedoosugar5rnutions. Fur,galspore<'ge1mfr1ate'quickly
Fa lse-colourSEMofyeast whe,evertheyoccu,
cellbudding(x4SOO}
= "~":,;;:'i::.:,.,.·:::-'•b1e
gromngcond,oons
mycel,umof
Mould fungi hyphae
feedooexposed, origina ls.pore
=~•e<gank
hyphaeandfungal
fruitingbodieson
thesurfaceo fthe
,w~"~ -~u
dung of he rbivores
'"""°"'""''
~~:d,spmed
typical succession
of fungi
Figure18.18TheFungi
435
m Bicxliversity. classific.ation and con~Mtion
• Multicellu lar organisms, differentiated into tissues; they are non-motile organisms (but the male
gametes d mosses and ferns are motile).
• Cells are eukaryotk, with a distincr nucleus and membrane-bound organelles.
• Nucleus contains linear chromosomes of DNA with histone protein anached.
e Lllrgeribosomes(80S)present.
• NutritionisautO{rophic .
• C..llwallsarelargelyofcellulose.
Figure18.19The?lant""
436
18.lClassif:ation I
Vertebrate an imals
437
m Bicxliversity. classific.ation and con~Mtion
Pos,lblaevolutlonaryralaUonshlps
batwHntha flva klngdoms l = orkjinoiile
Figure18.21 Possibleevolutioo~ryll'!al>omhipbetweenthefivek;<>goom,
Characteristics of viruses
Viruses are disease-musing agents, rather than 'organisms· . The distinctive features of viruses are:
• They are nOI cellular structures, but rather COflsist of a core of nucleic acid surrounded by a
proreincoat,calledacap.sid.
• In some viruses there is an additional external envelope of membrane made of lipids and
proreins,asinHN. Figure10.15,page204andtheinfluenzavirus)
438
18.3 ConseNation I
1 ;;~·= @
RNAviruse•
~ze:25nm
• They are extremdy small when compared with bacteria. Most
virusesareinasizerangeof2o-400nm(0.02--0.4nun). They
become visible only by means of the electron microscope.
""m~
immunodefkienc.y
viru,ol"'1imalhosts
$ • Theycanreproduceonlyinside specific living cells, so
virusesfunctionasendopara.sitesintheirhOiSCorganism.
• Th'-,Y have to be transported in some way between hosts.
• Virusesarehighlyspecifictoparticularhostspecies. some to
plant species, some to animal species and some to bacteria
2double-straooed • Virusesareclassifiedbythetypeofnucleicaddthey
reovirusof contain , either DNA or RNA. and whether they have a single
"'1inalhom
ordoublestrandofnucleicadd(Figure 18.22)
Figure18.22Thecla,m;c.ationofviruses
~ 18.3 Conservation
439
m Bicxliversity. classific.ation and con~Mtion
Stepsoffoss;lfonnatioobypetrilication
1. dead ,email"IS of organisms may fall in to a lake or sea and become buried in silt or sand,
inanaerobic,low-temperaturecondrtioos
3. rema;ns hardened in this way become compressed in layers, up thrust may btir,g rocks to
thesu,fac:e anderosK>noftheseroc:kscommences
4. landmOYementsmayexposesomefo,o;ilsandafewa, ediscove,edbychanc:eblll,of
the relatively few organisms f=il ised, ve,y few will ..,er be found by humar,<; Sedim e ntary rock layers and the rema ins of extinct
fossil species
Jn the long history of life on Earth, humans are very recent arrivals. Life origin3ted 3bout 3500
milHon years ago, but our own species 3rose only 3 little over 100000 years ago. Initially, human
activities had little impact on the environment. For one thing, during early human 'prehistory',
population numbers were low. Homosapi<ms were a rather 'struggling" species, living among many
verysuccessfulones.Atthisstage.survivalmusthavebeenaverychancyaffuir.
LookbacbatFigurel0.2onpagel94
The first significant increase in the human population occurred with the development of settled
agriculture - a change that began in the fenile crescent in the Middle East about 10000 years
ago. The current human population =plo.sion beg3n 3! about the beginning of the Indus trial
Revolutio n , 3round 200 years 3go, 3nd it continu~s today. No one is certain when the rate of
population growth will slow down, as it surdy must.
When we plot world human population against time we see a steeply-rising ]-shaped curve. Our
population growth 3ppears to be in an 'expon e ntia l (log) phase· o f growth. This is due to high
440
18.3 ConseNation I
binh rates and lower death rates, leading to rising life expectancy (people are living very much
longer). Tcxlay the impact of humans on the environment is very great indeed - there is virtually
no pan of the bioophere which has noc come under human influence and been changed to some
extent.Manyoftheseehangesthreatenthebiodiversityinallecruystems.
O nly a part of planer Earth, its land. oceans and atmosphere are inhabited. Living things inhabit
a narrow belt, from upper soil to the lower atmosphere. or, if marine, most occur near the ocean
11 Vvhatisthechief surface. This restricted zone is called the bios phere. The distribution patterns of living things
ecosystem in the within the biosphere show large. srnble vegetation zones. These zones are called ~'«>S}'Slems .
countrywhereyoulive7
Examples include tropical rainforest and grassland. Here communities typically extend unbroken
over thousands of square kilometres.
Thisbarchartisas.ummary
of the percentage
ofotgani,msidentilied
as"Critica llyendangered".
"Endangered'arid 'Vulnerable"
within several groups.Data
fiasbeentakenfromthe
IUCNRedli,twebsite
~=i~~: 2
d list.org)
J~-
Fora llgmup,othertflan
thoseabove.thelUCNha""
oolyassessedatinyproportioo
olthespecie,;nthosegmup,.
socomparison,canJlO!bem;,de
betweenandwithinthesegmup,
~~--~
Thethreeg,oup,oppmite
oow,,,.,,fia,,.,been
comprehensively assessed
-======i
andskates _..
Figure 18.24 1UCNRed
List data 0 5% 10%
441
m Bicxliversity. classific.ation and con~Mtion
442
18.3 ConseNation I
progressivdy less varfation in the population as the number of loci at which there are homozygous
alleles increases. Whilst the individuals of that population may initially be well adapted. in the face
of environmental changes they are less able to adapt. We can say that the 'genetic fitness' d the
population is compromised.
In the case of small, isolated populations, genetic variability is most critical. For =ample, in
populations of species that become endangered, it is a question whether they posses sufficient
geneticdiversitytobe ableroadapttofuturechangesintheenvirorunem. lfthey do not,their
survival is unlikely. This is a practical problem facing modern zoos attempting captive breeding
programmes (page 445). It is also a n issue for Ol:her attempts to protect endangered organisms
in the wild where population numbers have been reduced to small, isolated groups in former
strongholds
443
m Bicxliversity. classific.ation and con~Mtion
;,. o
functionedasasanctuaryforwildlife!
.,,. Today, this solution to extinction pressures on wildlife
includesthesettingupandmaintenanceofareasofspecial
scientificinterescasnaturereserves,ofnationalparks(thefirst
national park was Yellowstone, set up in North America in
. ... rt. . . .. 1872), and of the African game parks which have been more
recendyestablished. lntornl,thesesitesrepreseruhabitatsof
many different descriptions, in many countries around the
world. Some of the conservation work they achieve may be
carriedoucbyvolurueers.
Wbat b;ogeogruphicalfealuros ef a reserve best promote
0
pro!ect.
Also,thereis an edgeeff<--cl .Acompactreservewith
ineffective design
minimal perimeter is Jess effective than one with an =tensive
interface with its surroundings . lbe use of the surrounding
are-a is import:1m. too-, if it is managed sympathetically it may
••
indirecdysupponthereserve"s wildlife.
Another feature is g eographical L,;olatlon - reserves
•• •••
siruatedatagreaterdistancefrorna1herprotectedareasare
lesseffectivethanreservesthataredosertogether. Also.
it has been found that connc_-cdng corridors of land are
advantageous. lnagriculruralareasthesemaysimplymke
the fonn of hedgerows protected from contact w ith pesticide
Figure18.26featunesofeffectivenaturereserves tre-atmentsthatne3rbycropsreceive.
444
18.3 ConseNation I
The reserve or park isbestmanagedtotheadvamageofwildlifeby:
• the employment of local people as trained wardens and rangers, who patrol the area to ensure
people comply with local regulatiOfls. lbey noce the cOfldition of the wildlife, record the
changingpopulationsizesandprotectthewildlifewheretheneedarises
• the supervision of agricultural and hunting activities, mainrnining the emphasis on traditional
praaices applied to modest or minimal levels
• the control of permitted industrial aaivitie.s, typically mining or quarrying , logging, and fishing in
marine rese-rves, and of the associated transport infrastruaures needed.
Forvisitorsandecotourists
• visitor centres are provided and staffed to educate and inform Ofl the local issues of conservation
• access is controlled and restriaed to specific footpaths, tracks and roads to avoid critical wildlife
Continuousmonitoringofthen.-servesothatcausesofc:hangeareunde1'tood.changemaybeanticipated
and measures taken early enough to adjust rnndibom without disruptkm. shoukl this be necessary: fe<
exampte. rnral .._..,15 req uire monitoring for the imp,>ct of erntourism
Mainte na nce of e ffective boundaries and the limitatkm of unhelpful human interference . The enthusiastic
involvement of the local human community communicates the messages of the purposes of rnme,vation (a
local "educatkm" p,ogramme. in effect) and that ..,e,yone has a part to play in rnnservatkm
AJso. protectionofmajormigrati onroutesoflargeherdsofherbivoresandthebigpredatorsthatfollow
them.asinAfricaandNorthAmerica
14 1/v'hatarethe Measurestofaci li tate thesuccessful completionoflifecydesofanyendangeredspeciesf0<\\+.ichthe
particular challenges in rese,veishome.togethe,withsupportiveconditionsfmvulnerabteandmespecies
the management of a
Restocking a ndreintroductionsofoncecommonspeciesfromstockspmducedbycaptivebteeding
nature reserve in your
programmesatzoologicalandbotanicalganlens. Thisma~espo,sibjereintroductionofspecies(us.ually
locality or country?
predators) hunted almost to extinctkm in farming areas. including wolves in furope and in the northern
nationalparksintheUSA
445
m Bicxliversity. classific.ation and con~Mtion
Figure 18.27 Welkome Trust Millennium 8uikJing. Wakehum Place name to the Millenn ium Seed !lank (an
initi.ativeoftheRoyal8otanic:Gardens.Kew,intheUK}
446
18.3 ConseNation I
New developments in conservation
Using assisted reproduction
lhe introduction of capiive breeding programmes was discus.sed on page 445. Techniques developed
in modem reproduaivc technology have been adapied for use in these programmes (Figure 18.28).
Representative endangered species of mammals, held in zoos, may be effectively conserved by
theprocessoffertilisationofeggsoutsidethebody(h1.vhrofertilisa lion-
IVF). The key srep in TVF is the successful removal of sufficient eggs from the ovaries. To achieve
this. nom,a] menstrual activity is temporarily suspended with hormone-based drugs
Then the ovaries are induced to produce a large number of eggs simultaneously, at a time
controlled by the team of zoo vets. In this way the correct moment to collect the eggs can be
known accurately.
Eggcellsarethenisolated
from surrounding follicle cells
andrnixedwithspenn.Viable
sperms may be held in sperm
banksinafrozenstate,and
therefore can be from male
animalsheldinzooselsewhere
in the world. lnthisway,the
gene pool of endangered
spedesissignificarulyles.s
restricted.
lffertilisationoccurs,the
fertilisedeggcellsareincubated
sothatemb.-yosatthe8-cell
stagemaybeplacedinthe
uteru.s.Jfone(ormore)imbed
there.thenafX}l"malpregnancy
may follow. Spare embryos can
alsobefrozen,andtheyroo
canbe.sharedwithO(herzoos
The young mammals created
inthiswaymaybeexchanged
too,andsurrogatemO!hers
may be used. The steps to
IVFasiJJustratedareroutine
inprindple,butinmany
endangered species the success
ratehasbeenlimi!ed,sofar.
JM,aoscopk.....,..natlootocooflrmtllaternblyoshaYe
f&IChedlhe4-to!kelstage
Figure 18.28 The pmce,;s of in·ITTm fertilisation, adapted for animal cofl'ieivation
447
m Bicxliversity. classific.ation and con~Mtion
Figu~18.30Contracept,Ve
b~itdepk,yedfor tf>e grey squirrel
populati0<>
448
18.3 ConseNation I
The rabbit- a biological control measure
Anocher example of an alien species that has become 'invasive" is the
rabbit (Ory:ctolag,,s sp.), which was deliberately introduced from Europe
into Auscrali3 in 1859. This voracious herbivore spread rapidly. No natural
predator5 present were an effective threat (inle ..,. pecifie com petition failed),
:mdrabbitsquicklyoverranlargeparuofthecontinent. Grassl:md, available
to herds of herbivores, principally cows and sheep, was seriously damaged.
Agricultural produai= was plunged into a crisis. This continues. even now,
withvaryingdegreesofscverity.
In desperation, and as a bio logic al con1rol m easure , myxoma virus that
had been discovered in South American rabbits (a different species from the
European rabbit). was introduced in 1950. Myxoma virus causes a parasitic
disease of rabbits, myxamatosls. but on its "home" continent its effects were
mild. lbe changing effects d this virus Ofl rabbit monality (1950-19S6) are
shown in Figure 18.Jl, as is the resulting change in virulence of the virus. We
Figure18.31Char,gir,grabbitmortalityandvirus may assume that initially. a small number of the huge populatiOfl of rabbits
v1rulenc:e.following;n1roductionofthemyxomavirusasa
biologi::alrnntrolmeasure had immunity. As their vulnerable relatives were kilJed off the immune rabbits
prospered from the diminished competition for grass. Rapidly, the bulk of
the population were immune, and any that failed to develop immunity were
quickly taken out.
lntheiroriginalhabitatan'alien'specieswUlhaveevolvedinthecompany
of natural par:,sltes and prcdatot"!! - and come to =ls! in balance with other
organisms of the community. In a new habilat the alien"s natural enemies may
be absent. so thry grow at the expense of native species. crowding them out.
Japanese knotweed
A most unfonunate example of this is Japanese
knotweed (Fallopiajaponica), deliberately
introducedintonorthernEuropeintheearly
nineteenth century as an ornamental plant for
garden ponds and Jakes.
TheadvanceofJapaneseknotweedinthe
UK is an =ample of the spread of an invasive
speciesdevoidoftheusuallimitingfaaor:sthat
keep it under control in its native Japan (Figure
18.32). Here, knocweed plants grow into dense,
submerged thickets that overshadow and crowd out
nativewaterplants,blockwarerwaysandpublic
acressroscreambanks.lnJapan,theplantgrows
inabalancedrelationshipwithotherwaterplants
and causes no significant problems. Meanwhile,
in Britain, the chemicals in the plant"s leaves and
rootsdiscouragepredmionbylocalleafbrowsers
or root parasites. In Japan, predators have evolved
thatcancopewiththesedefencemechanisms.
Farmers, gardeners and local councils face huge and
increasingbillsrosuppressthisrampantinvasive
species.
Figure18.32Distrtilltionofknotweed;ntheUK.1'l00-94
449
m Bicxliversity. classific.ation and con~Mtion
You ran read more about the work o f the WWF at: www.worklwildlifc.o r g .
45-0
18.3 ConseNation I
the initial disaster, removal d oil slicks may be speeded by spraying the oU with essential inorganic
nutrients,chieflyphosphatesandnitrates,whiehaidthesaproi:rophicbac:1eriaandthu.sspeedup
baaeriologimloxidation.
Figure18.lllrwestigatir,gthe iooc:ulumoioilydirtscra,ped
deg,adabonofmmeraloil",p;lls" fromc:oolerpa<tofc:¥eogine
(shakenupwrthoilaJ1dnu!rieJ\ts
atstar!ofexpe,iment)
- weighedsample7
Timir,goftr>enumbe,olho<xs
;~~1~~tionand
s.ma l volume
ofeogineoll
(known volume)
essemial minernls
(known volume)
Bioremediation of soils
Pcstld d cs are prcxlucts of the agrochemical industry that are widely used ro control harmful
org:misms thm are a danger to crops or herds. Their use =tends to the honkulrural industries
tha! arenot 'organirnlly'orienta!ed.ofcourse,andtheuseofpestiddeshasbeenenthusiastically
adopced by many people who maintain gardens in developed countries
Mcxlern pesticides are substances de-signed to kiJJ specific types of pest, including plant weeds
(by herbicides), insects(byinseaicides),fungi(byfungicides). andslugsandsnaUs(molluscicides).
Pesticides have enonnously improved produaivity in agriculture, but their use has generated
problems in the environment. Herbicides are the m06! widely used group o f pesticides, but
insecticides are also widelr used and they may cause the greater problems for humans.
Pesticides are a wide range d different compounds. Many of them are organic molecules
suitableasrnrbon sourcesandelCC!rondonorsforvarioussoilmicroorganisms,but othersare
not. Those that can be bicxlegraded are eventually removed from soils (fable 18.6), but their
breakdown may not be entirely due to the activity of microorganisms. Some substances leach away
into underground water, and others may spontaneously degrade with time.
Table18.6Persi,tenceofa
ITimefordisappearance fromsoil
selectiooofpeshc:ides
2.4· D l 4wpek,
l 20weeks
lnse cticide(chlorinatedcompound)
l 4ye,m
lnsecticide(organophosphatecompound)
Malathion I lwl'ek
l lwl'ek
451
m Bicxliversity. classific.ation and con~Mtion
lnd n strlal soh "ents, now widely used in devdoped court{Jies at le3st, are organic compounds
such as dichloroethylene, trichloroethylene, chloroform and some brominated 3nd fluorinated
related compounds, to name but 3 few. These toxic subst:mces (some are also suspected of being
carcinogens) are frequently detected as persistent conrnminants of ground waters, This is 3 wdl-
documemed problem in p:,ns of the USA, for example,
A variety of different bacteria 3re 3ble to bre:ik down these molecules, including the removal
of the chlorine component, For example, the bacterium Debalobacterium uses the compound
dichloromethane in its metabolism, Substrate-level phosphorylmion (ADP + P; ----> ATP) occurs in the
Jn areas where the underground W3ters are seriously polluted, the activities of natural
populations of bioremedi3l microorganisms m3y be enh3nced in silu by the drilling of wells and
vents so that micronutrients 3ncJ sources of oxygen (as compressed air or peroxide solutions) can
be 3dded for the benefit of the microorganisms, 1be natural de:in-up reactions are 3creJerated in
this way, although the process may rnke months 3nd years to complete, No doubt the dispersal of
thepollurnnts3ndthedepthofthegeologicalstrarntheyoccupyis3factor,
e lheprocessofclassif1cationinvolvesnamingorganisms differentiatedintotissues.Manyaremotileorganisms.Their
usingthe bin o mi a l system sothateac:hhasageneric nutritionisheterotrophic
andaspecificname.Generaarearrangedinahierarc:hical • Biod ive rsityreferstothe vastnumberoflivingthingsthat
daslification, of which the kingdo m is the largest and mo5t exist. About 1.7 million different species are known, but a
indusivecategory. There are five kingdoms muchlargernumberoforgani:smsmayexist,undiscovered
e The Pro karyotae{prokaryotes)includethebacteria, The needto ma inta indi versityisbasedonecologic.al
cyanobacteria(photosyntheticforms}and archaebacteria arguments,but in dudesdiversityasthesourceofgenesfor
{extremophiles}. They are unicdlular or filamentous improvement of agricultural crops, as a source of new drugs,
prokaryotic organisms consisting of very small cells, 1~5µm toresistclimatechange,tosupportecotourismandforethical
Theirnutritioni:sheterotrophicorautotrophic and aesthetic reasons
e The Protoc.ti st a{protoctists)includethealgae{indudingthe • A spe<ies is a group of individuals of common ancestry that
multicellular seaweeds), the protozoa and the slime moulds. dosely resemble each other and which are normally capable of
Theyaresingle-c:elledeukaryoticorgani:smsor,ifmulticellular, interbreedingtoproducefertileoffspring
notdifferentiatedintotissues.Theirnutritionisheterotrophic • The exponential expansion of the human population is a
orautotrophic relatively recent phenomenon, but the t urre nt size o f
• The Fung i include the moulds, yeasts, mushrooms and th e human po pula tion results in huge demands on the
brac:ketfungi. Theyaremulticellulareukaryoticorganisms Earth'slivingspaceandresources.Manye nviro nm e nts are
(excepttheunicellularyeasts),notdifferentiatedintotissues be ing deg ra ded and biodiversity is being reduced. Today,
and non-motile. Theirnutritionishetemtrophic. speciesareindangerofex tin t tion duetothelossoftheir
e The Pl a ntae{thegreenplants)includesthemosses,ferm, naturalhabitats,changesintheenvironmentordueto
ronifersandfloweringplants{broad-leavedandthegrasses} over-exploitation.Thesecausesareallattributabletohuman
Theyaremulticellulareukaryoticorganisms,differentiatedinto activity.
tissues. The plants are non-motile organisms, but the male • Co nse rva tion isthemanagementofresourcesandhabitats
gametes of mosses and ferns are motile. Their nutrition is soasmaintaintheexistingrangeofhabitatsandtoatternpt
autotrophic topreventluturereductionsinbiodiversity.Thisindudes
• The Anima lia (Animals) includes the non-vertebrates, such as ca ptive breeding prog ra mm esandtheestablishmentand
worms and arthropods (which includes the insects), and the maintenanceof rese rvesa ndna ti o na l pa rks,wherethereis
vertebrates,whichincludethefish,amphibians,reptiles,birds supervi5edprotectionandanopportunityforpeopletoleam
and mammals. They are multicellular eukaryotic organisms, about biodivenity and its conservation
452
Exar::'oationstyleq..,stions .
buttresses at
baSeoltrurik
-
)O<lngtree
t o Iii a gap
Fi g.1.1
2 TheAfricanelephantooc:eroamedmo5tofthecontinentof
Africa. The table below gives data on more recent estimates
of the population sizeofthisgiantherbivore
Population size
By1979
By1989 600000{<1%0/original
numbers)
a) Outlinethediverserangeoflactorsthataremostlikely
tohavedirectlycOfltributedtothisdecline [12]
b)lfherdsoftheAfrK".ilnelephantcontinuetodeclineshilrply,
explainwhatimpactthisislikelytohaveonheterozygosity
intheseanimalsandthe impactthischangeislikelytohave
[8[
[Total 20]
4Sl
A Level
19 Gene technology
The discoveryoft hestructureofDNAbyWatson a nd Cr ickinthe structureandprotein synth es isstudied intheto piconNucleic
ea rl y 1950sanddi scove riessince haveled to ma nyapplica tion s acid sa ndprote in synthesis. W he reposs ible,studentsshould cury
o fge ne t ec hno logyin areasofmed icin e.agricultu rea ndfo rensic o utpra ctical w orkusing e lectropho res is,e ith erwithDNAor
sc ie nce.Thistop ic re lies heavilyo npriorknowledg eofDNA specia lly pre pared dyes used torep resentDNA or proteins.
Genetic engineering By the end of t his section you should be able to:
involves the manipulation
of naturally occurring a) define the t e rm recombinant DNA
processes and enzymes b) explain th at genetic engin,,.,ring involvoes t he extraction of genoes from one organ ism, or the
Genomesequeoc:ing synt h"'isofgenoes,inorde rtoplacethe min a nothe rorganism(ofthe same oranotherspecies)
gives information about suchthatthe re ce ivingorganism exproessoesth e ge ne product
thekx:ationofgenesand c) doescr ibe the p rinciples of t he polyme rase chain reaction (PCR) to done a nd amplify DNA (the role
provides evidence for the ofTaqpolyme rase shouldbeemphasise d)
evolutionarylinbbetween d) doescribe and expl a inhowg e l e lectrophoresisisu,edtoanalyseproteinsandnud e icac:ids,andto
d istinguishbe twee nthe allele,of a ge ne
e ) doescribe the properti"' of plasmids that allow them to be used in gen e cloning
f) explainwhypromote rsandothercontrol,eque ncoes mayhavetobetransfe rredaswellasthe
doesire dgene
g) explainthe use ofgenesforfluore ocentor easilyrtainedsubstances asmarkers inge ne technology
h) explainthe role,ofrestriction e ndon udeasoes,reverse transc:riptase andl igasesinge netk
engin,,.,ring
i) explain, in outline, how mkroarrays are used in the analysis of ge nom"' and in de tecting mRNA
instudi"' ofge neexpr"'sion
454
19.1 Principlesofgenetictechnology I
The techniques of the geneiic engineer offer numerous practical benefits in biocechnology.
medicine and agriculture, and also to science and the study of how genes operate in the COfltrol of
cells of organisms. At the same time, outcomes of geneiic engineering may generate environmenrnl
and c thk al issrn.·s of concern ro society. We will return to this issue later.
Other applications
Today, in addition to the cre3tion of GM organism.,, gene technologies are applied to:
• DNA st.-qucncin g - the creation of genomic libraries of the precise sequence of nucleotides in
samples of DNA of individual organisms. The nucleotide sequence in the whole human genome
was the product of the Human Genome Project. Today, ocher genomes have been completely
sequenced, including those of some viruses, bacteria, fungi and many ocher eukaryoces.
• Gc n <-'t k fingerprinling - in which DNA is analysed in order to identify the individual from
which the DNA was taken to establish idemiry and the genetic relatedness of individuals . It is
fX>W commonly used in forensic science (for example to identify someone from a sample of
blood or other body fluid). It is also used to determine whether individuals d endangered
spedesincap1iviryhavebeenbredorcap1uredinthewild.
Table19.1Thegenetictec:hnok,gist'stoolkitolenzymes
These enzymes a,e extriKted mainly from mkroorganimis Of viruses and are used to manipulate ntKleic .Kids in
1 gening a ropy ol
theqer,etobe
eogir>ee<l'd
2gene intovector
lortransporl to
newagan,sm
Figure19.1The1tepsinthegeneticengineeringoff.ro/ilorimulinproduction
--
S switchir,goo
455
m Gene technology
purif>edmRNA
=l
codir,grin,ulin
synthesisofcDNA
aw"'"""""'""'""~""''-'"°"'""""''
romplementary<lrandofDNA (usingapoololmdeotides)
bybasepai"ingwi!htheseq.,..oceolbasesofthemRNA
mRNAbeingusedasatemp!ate
lorDNA'Yflthesis
cDNAstrand
- DNArnmpjementa,yto
b.asesequeoceofmRNA
DNA polymerase
synthesisff
~N":!~:~~tary DNA polymerase
en,ymei>dded -
i m;,,yn1he<isesasecond
DNA strand.complementary
'T
copy of human
=baseseqlll'IICf'of
456
19.1 Principlesofgenetictechnology I
Once the gene is isolated, the double-stranded DNA undergoes amplification; many copies are
made. There are various situations in which a genetic engineer is able to produce or recover only
a very small amount of DNA (such as at a crime scene). It is now possible to submit such minute
DNA samples to a process known as the polymcra.sc c hain rcac1ion (PCR). In the polymerase
chain re3ction, DNA is replicated in an emirdy automated process, in dtro, to produce a large
number of copies. A single molecule is suffidem as the srnning mmerial, should this be all thm is
available. lbe product, are always =act copies. The heat-resistant polymerase enzyme used in this
process is known as T<1q polymerase after the thermophiHc bacterium. Tbermo,s aquaticus, found
in hot springs and hydrot:h.,,-mal vents, from which it was originally isolated. This pru1ein is able to
remain an effective catalyst at a temperature of97.5°C for the length of time required in the PCR.
5' knownendsequeoces r
1 0ouble...trandedONA\\ith
2Separatechaimby/J'
heat,ngto95°C {
5' 3'
3Makeprime rthatmatchesthe
endsequencesoftheONA
6Repeatprocess
fragmenttobecopted usingthetwonew
Coolto40°Ctoallowthe
double strands
prime,tosticktoeachstrand
3· TIITT 5·
4~d~;~~;~~~-:. . 3'
'>--r~r5'
ntKk,otides, and 5 Nudeotidesaddedtoprimeisusing
heatto72"C singlestrandasteJr4Jlate
Note: 'Primeis'a reshortsequeocesolsingle-strandedDWl.madesyntheicallywithbase
sequeoces com~imentary to one end {the 3' end) of ONA
Remember. OWi. polymerase synthesises a ONA strand in the 3' to 5' di"-'CIKlO
Figure19.3Thepo/ymerasecha in,eaction
457
m Gene technology
~ isolatedplasmklo,o•
~
base sequences in
!1 1!1!1 11 1)1111! ONAandC\ltsthe
doubH,helixatthese
cTrt c!t ("TT AA
6 particu!a,points.
leavingstidyends
ONA strand cut by plasmidC\ltby cLi
restfictOOOenzyme :\~~~el
!
AATT
thatrecogniSl'S the
seq uenc:eGMTTC
"stid<:yend"
,..............
sequenceGMTTC
~ " b d y h t t~1 ~
~
TTTTIIIIIIu..u
~nd TTAA
' A; ....,.-'
"stidy
: 7-:.~~-
-._ base pairing ocC\lrsat/ end"
Splici ng mear,sthe le id ds
h4rr Ligasejoinsthesugar·
phosphatebadbones
ofthepairedstickyends
ligasecatalyses
formation of a
C--Obondhe,e
Figure 19.4Genesplicing-theroleofthell'Stfictiooenzymeandligase
Stickyendswiththesamesequenceasthoseontheplasmidarealsocreatedontheisolatedinsulin
gene. This is done by adding shon lengths of DNA that are then 'trimmed" with the same restriction
enzyme. Jn this way complementary "sticky ends" now exist at the free ends of the cut plasmids
and the cDNA of the insulin gene, making it possible for them to be 'spliced" together into one
continuous ring of DNA. The enzyme ligase catalyses the formation of a new C--0 bond between
ribose and phosphate of the DNA backbone5 being joined together. Energy frcm ATP is needed to
bring the reaction about. The steps are summarised in Figure 19.5
458
19.1 Principlesofgenetictechnology I
A note on restriction enzymes
Restriction enzymes occur naturally in bacteria. They are believed to have evolved as a defence
mechanism against invading viruses because inside the bacterial cell they selectively cut up foreign
DNA. Many different restriction enzymes have been identified as useful by genetic engineers . 1hey
are named after the bacterium frcm which they a"' isolated (genus, species and scrain).
0'
p!asm,d
a ,estrictOOO enzyme
ruttingDNA,e.g.at
~
to the rut plasmid open plasmfds alld
1
groeffagmenlshave theenzymel,gase
;ftc•"
recomb,nant plasm,d )
MTIC
G
.:~:~<k ~TIM NNGAA L
NNC TTAAGNNNN
CNNNN
Figure19.5Usingaplasmidasthevecto,/ortheinsulingene
459
m Gene technology
tempora,ycreaoonoi
~nyporesin iooooo
walsoino<tbacteria,
aNowfr-,georryof 00 00 0
r«ombina nt
plasmids ,ecomb<l1amplasmids 0 0 0 O
0 O O
..J.,~.,,
Figure 19.6 The,etumolrl"Combinantplasmidstotnl'bacter;um
460
19.1 Principlesofgenetictechnology I
Finally, whenthetwopolypepcidechainsareseparntedfromthebacterfatheyaretreatedfirstwith
cyanogen bromide to cut them free from the longer polypqxide chain at the methionine amino
acid. Then they are funher treated to promcxe disulfide bond formation and development of a
spec,alplasmidwi!hgeneslor insuingme'
ll'sis!anct'to tetracy<line,aod
antibioticresirtana - ooegenegi,,;ng
+manyun<hang,ed
gene/or plasrmh
amJ)ki lin
~:~:~~;;,·::i.':"'i:::.·
willgmw.Fromthis mastu platotreplic:aplates
~ - - - - a,emadeusingaslefile'replka platingpad' - - - -~
(shoNnbelow)
thebactPriaol!hiscolooy
f.;ledrogmwonplatell,
~:100\m::i~·:,,:: ~ •
• •
•• • •
•rI••• •: ---------- .. .• ••.•
•
thesebacteriaareoowrnltuf<'dinthe
laboraroryroconfirm!heyprodoce
insul:n(finalse4ectK>llsiep).prio<ro
rnlruringinaniod,mriallermemer
{commercial insulin manufactu re)
461
m Gene technology
• the transcripiion of eukaryotk DNA to messenger RNA is more complex than in prokaryotes: it
involves the removal of shon lengths of ·non-infonnative" DNA sequences, called ' introns"
5 Whyarethegenesof
prokaryotesgenerally • the mechanism for triggering gene expression in eukaryotes is more compl= than in
ea1iertomodifythan prokaryotes. Genes for the promoier and other control sequences have to be transferred. as
those of eukaryotes? wellasthedesiredgene
Despite these difficulties. severalvarietiesoftransgenkplantsandanimalshavebeenprcxluced.
462
19.1 Principlesofgenetictechnology I
possibility must exist. 1be outcome would be an 'engin=red' dise'llse-causing microorgani.sm that
Genesforfluorescent(oreasilysmined)sub.stancesarenowrn.edasmarkers(Figure 19.8)
in.stead of antibiotic resisrnnt genes, alongside the cDNA of the desired gene. These genes are then
linked to a special promoter. The marker gene is =pressed only when the desired gene has been
successfully inserted into the genome of the host. Org:misms that have been transformed in this
way will glow under UV light (or can be identified by staining)
Thecrysta lje llyfish(A equorea victoria) 0« ursinthe Fluo resce nce is du e tothe presenceof
planktonoffthewest coastofNorthAmerica. ltsn<>W'i ag reenfluoresce ntp rote in(GFP). This
bright greenflumescencewhenexposedtol>ght;ntheblue proteinandthegenethatcodesforithave
toultravioletrar,ge beenextr.tcted
Thetertia,ystroctun,ofG f'Pprote;n
Th e GFP ge11e is now used Ha marker The GFPgene hasbeenintroduce d andm aintainedin
gene in genetic engi neering. ~Of th ege nome,andthe n expressedin spe ci esofbacteri a.
example,itmaybeirisertedintoplasmid, yeast, fis h. plants. insects. and mammals. f'or example.
ak,ogsideaclonedgeoeandilspmmoter. tfiese GM mice glow green under blue light because the GR'
When thesepla,mKlsare,etu,nedto genehasbeen;ntroduced intothe;,DNA.
bacteriaandthepmmote, isactivated, the
transformedwcteriawill bedetectedby
expmuretobluel"]ht
--~ "'""'
GfP9"""'
promoter
Theteniarymoctureo!GfPproU.n
Figure 19.8Theuseofffooresc:entmari;e"
463
m Gene technology
As a result af the llunmn Genome Project (HGP), the location of each human gene and the
base sequence within its DNA structure are known. Now, where a single--stranded DNA probe,
complementary ro a panicular gene can be produced and made radioactive or fluorescent in
ultraviolet light, original genes can be located and isolated. To do so, the DNA has first to be cut
imo fragments. Fragments are then denatured into single Slrands by heating, separated by means
of electrophoresis (Figure 19.9), and then treated with the probe. The fragments of DNA conmining
the gene are located.
..-,
Bothofthesemethodsarecurrentlyapplin:ltoisolategenesrequiredforothergenetechnology
Electrophoresis
Ekctrophon,sis is a process used to separate molecules such as proteins and fragmems of nucleic
acids. It is widely applied in many studies of DNA. For example, it is central to the investigation d
the sequence of bases in panicular lengths of DNA, known as DNA s eque ncing. 1t is also used in
the identification d individual organisms and species known as genetic fingerprlming. We will
review these applications shortly.
464
19.1 Principlesofgenetictechnology I
• l•ctrophoru isinprog,..,
Figure19.9Electrop00.esis-theseparat;oo
of DNA fragments
DNA microarrays
DNA microarrays are small, solid suppons, typically made of glass, silicon
or nylon, each about the size of a microscope slide. Omo this surface, DNA
probes that correspond to thousands of differern genes are attached in a regular
panem of rows and columns. Each dOI on the array conrnins a DNA sequence
that is unique to a given gene. 1be microarrny is then used to measure changes
inexpressionlevelsofindividualgenesfromcellsunderinvestigation.
The DNA that is to be tested originates as mRNA being =pressed by a
panicular cell. By the action of reverse transcriptase this mRNA is convened
to cDNA. As the cDNA is being synthesised it is simulrnneously linked to
fluorescent dye. Then the DNA probes of the microarray are =posed to this
cDNA. Any o:implementary sequences present will bind ro the fixed probes
After this. the microarray is rinsed fr.,._.. of any cDNA that has nOI hybridised
and become bound. Finally. the microarray is exposed to laser light that causes
fluorescence where there has been hybridisation between cDNA and a DNA
probe,andthere:sultsrecorded. Thebrighterthelighteminedthehigherthe
levelof=pressionofapaniculargeneinthecellfromwhichtheoriginal
Figure19.10Mic:ro;mayanalys.i,mac:htnewith
mic:m,maybeireg;nserted
465
m Gene technology
~ ~
J M~<mS,, J
-.r'V"- .r mRNA ::;:--v::;:
~ ~
eac:hspotolDNA
teprl'Sl'f\tsa
o:fferef\\gelll'
Figure19.11M>Clo.amyanalj,,;i,ofdifferential91'11eexp,es,io11
466
19.l Genetic techno~gy applied to medicine I
19.2 Genetic technology applied
to medicine
Bioinformatics
Bioinformatics is 3 new discipline that combines thr= major components:
• gt.·n o m ks - the mapping of genomes
• prot<.-om k s - the 3ru,lysis of the entire sets of proteins expressed by genomes
• b lo t<·drno loi,,y - the industrial and commercial application of biological science, particularly of
microbiology, enzymology and genetics
Developments in bioinformatics have created powerful t=hniques that allow scientists to find
similarities in nucleot:ide sequences and discover the function of proteins among organisms of
similar or different species. Nowadays, data is collected 3round the globe and from differeru
research groups. The purpose of the resulting databases is to facilitate the sh3ring of this
information, and ro allow the wide:st possible =ploitation of a huge and ever-growing body
ofknowledgeandskiJJs.
A flavour of the scale of developments can be gleamed from the following link to the 180
completegefX}fl}essequenced:
www.gcn o m c n cwsn c twork .org/n:sourccs/scqucn ccd_gcn o m cs/gcn o m c_guidc_p l .sh tml
467
m Gene technology
~ damagedbloodvessel
exposedcrn1<>genfibr._.,
, /~ anddamagedrodothel'-lm ~
,eieasedottingfacro,
(e.g.lac:to<Vll)from
SEMofblcoddot damagedtiswe
showingmesl"M'orkof
:::t!:'andna?J)"d
coomictioooltiamagedvessel ~ do!
stabiisedplug
· ~tsfurlhe,blood~,
" "'"'"''mentryofbaclefia
figure19.12Theblood..cJottir,gmechanism
468
19.l Genetic techno~gy applied to medicine I
Seven, combine d inununodcfidcn cy (SCID) is an inherited disorder due to 3bsent or
malfunakoing T- and B-lymphocytes of the immune system. The result is a defeaive antibody
response. In SCID patients the immune system is so compromised it is effeaively :,bsent, and
victims are extremely vulnerable to infectious disease from a very early age.
One form of SQD is due to a l3ck of the enzyme :,d c n oslnc d camin :,sc (A D), coded for by
a gene on chromosome 20. In the presence of a defective form of the gene. the substrate of the
enzyme this gene codes for accumulates in cells. Immature cells of the immune system 3re sensitive
totheroxiceffeasofthissubstratemolecule,3ndtheyfailtom3ture.
Now stem cells. h3rvested from the umbilical cord blood at birth. have been genetically modified
with the normal human AD gene (cDNA). introduced by retovirus. These GM stem cells are then
transfu.sedintotheneonateforsuccessfulgenetherapy.
So, in summary, gene therapy is an application of genetic engineering with the aim of supplying
a missing gene to body cells in such a way that it rem3ins permanently functional, when this is
thoughtsafeandethicallysound.Genethernpyisaveryrecent. andhighlyexperimental,science.
The steps ro the process of gene therapy are discussed below.
469
m Gene technology
~'!:::,~':',:;"..,~
"
d,.anneltofunctioo
AW
ADP+P ;
actWatiooeveo! -
themleolATl' m
-r - r
HowCFTRregu lateswater co ntfll'ltinmut us
·=·. ,. --r-
epithelium -
!issuefl uid -
___
Na•
c1-
waier
---
Na•
ci--
wale,
----
Na•
wate<
Figun,19.13TheCfTRprotein-achannelprotein
470
19.l Genetic techno~gy applied to medicine I
The process of genetic screening
We can now locate genes responsible for human genetic diseases, if we choose. This is one
outcome of the Human Genome Project. For example, the gene that codes for cystic fibrosis (CFTR)
was sequenced in 1989. This m:,de possible the screening of human cells for the presence of the
murated CFTR gene (Figure 19.14). So it is possible for couples robe genetically screened to assess
how likely tOC,y are ro have children who will have cystic fibrosis (because bol:h are 'carriers'), for
example. Alternatively, it is possible for all members of a family group be offered screening where
thereisahistoryofaparticularcondition.
Pr~-natal scree ning is used ro test for the presence of a number of genetic conditions in the
unborn child. This allows the detection of:
• chromosomes abnormalities such as Down syndrome (trisomy 21). trisomy 13 and trisomy 18
In the latter two abnonnalities. only very rarely do affected infants survive their first year.
• single gene disorders. such as haemophilia, sickle cell anaemia and cystic fibrosis .
• neural tube defects, such as spina bifida and anencephaly.
Prenaralscreeningmaybecarriedoutby:
• chorionk villu s s ampling- undertaken at ,veeks 11- 13 of the pregnancy, a sample from the
placenra is taken.
• amnioccnlcs ls - undertaken at weeks 15---20 of the pregnancy, the feta[ cells in the amniotic
DNA
1 -
2 DNAfragments separatedbyelectrophoresis
:~;:::':1~~- enzyme
fr,cubatedwilhresmctionenzyme - - - - --
,est,iction ·nm"atap.nirular
seqoenc:eofbase,
negati,,..char~ooDNA(dueto~1egroups)cau=
thelra9ffll'Jl!S !Omo"" to thepo,~M'electrode(anode).
bY1!he90lhasa"sieWl(j"effect - ,maHe,fragmemsmo""
mo,erapidlythanlarge,ones Figire19.14Stepsinvolvedinge!ll'ticscreeninglorcysticfibrosis
471
m Gene technology
, ho rioni<villussamp ling -
wilhdrawalofasampk,of!hefe!.11
!issuepan-buriedin tnewal lofth,e
lllerus iotheperio<l1l - 13week,
in!O !hep<egnancy;theti"fsample
i,of,el!s!hatareactivelydi,,;ding d,oriooic:
andu>nbeanalysedqo,iddy. viii
a mn·o,enteSs -
witrnlrawalolasampjeol
amnimkMuidintne"'°riod
:t~:=~;:~o:,,
tnesurtac:eoftheemb<yo
Figure19.15P"-'flatalscreening
472
19.l Genetic techno~gy applied to medicine I
The outcome of screening - genetic counselling
lbe outcome of genetic screening frequently does nOl necessarily allay a person's (or their pan-
ner's) fears. lbey are likely ro be referred to a gen e tic counsellor. The role of the counsellor
is to rnke a detailed case hisiory of the person and their immediate family. A counsellor may
thenresearchthefamilypcdigrcc.
Figure19.16Ageneticco,mselling,.,.;on
473
m Gene technology
'ey
Qoormalfemale o l>Offll,)1ma1e
@;::;female · =~mac
Figure19.17QueeoVlctoriaandherhei"-acelebratedcasel
474
19.l Genetic techno~gy applied to medicine I
4 Gene therapy
Somatic gene therapy applied to cystic fibrosis
Since the healthy CFlR gene codes for a protein that funaions as an ion pump, the gene therapy
initiative has involved getting copies of the healthy gene to the cells of the Jung epithelium. This
canbeanemp1edinanurnberofways.
The CFlR gene may be delivered in an aerosol spray. The spray contains tiny lipid bilayer
droplets called liposomes to which copies d the healthy gene are attached. The Hpooome.s fuse
with cell membrane lipid and deliver the gene into the epithelium cell (Figure 15.16). In trfals.
the treatment is effective but the epithelium cells are routinely replaced so the beneficial effects
only last a few days and the treatment has to be regularly repeated. lbe cure can only be more
permanent when it is mrgeted on the cells that make epithelium cells and unfonunately this
approachhasfailedtodeliverthehealthygeneeffectively.
An alternative approach is based on a viral delivery of the healthy CFrR gene. Viruses such
as the Adenoviruses have been used in place of liposomes. 1bis virus is capable of entering and
infecting Jung epithelium cells. However. the virus is first genetically engineered ro inactivate or
remove from its genc.ne the alleles that cause disease. Additionally. healthy CF!R alleles are added
in. Unfottunatclythecreatmentha.snotbeensuccessful. Viralvectorshavetriggeredallerg icor
other immune responses and have had to be discontinued
live, (ar,ytissue,emoved
will be regenerated) 1 portionoflive,is
- - - ,emovedfromthebody~
2brnkeninto
liverviablood,1... m Ga
indivodualce lls
~"'"'"'"'" ~
healthygeneadded
~ g,~ ~ /
4,evern>traflSCriptasegene,atesDNA.
whic:hisi>ddedtochromosome
*
3healthygene
addedintoretfovirus
Figure19.18Genetherapyusingav,msasvectot
• Naked DNA. Genes have also been transferred by direct take-up of DNA from the environmem.
lbe host cells have to be capable of taking up the DNA - a condition described as 'competent'.
Calcium chloride treatment has proved ro be one of the best methods of preparing competent
cells . The genetic diseases created in this way include SQD
475
m Gene technology
1 (FrRgenei,rntandadded!o o
~plasmkl,us;nga remictioo
e11z;mea11dthenaligase
I
/ bacteria lplasmid
I
I
I
cystic fib rosisgeJ>e,fuoctiooally
atransmembraneregularor,so
I
koownas !h.eUTRgene
', .0 /='•"""'
bacte,rialplasmid
/// (negiltivelyd>arged)
liposome
(enlarged)
liposom a-ONA
complex ///
---- ~ ~
----:=. ~~
2 liposomes,tiny li pidmembraoespherP<(positively
char~. rombiJle wit h DNA to facilitate emry
imolunga.>11,acrossthecel ,urlacememb<ane
4.1 liposome--ONAcomplexl=with
epW>eliumce l membrallE'
4.4 CFTRmRNApassestoribosomes
r'..l--t-~ incytoplasmandtheCTTR-coded
proteinisfo,med
In re ce nt clini cal trials ,ome 20% of epithe~um ce11s of CF patoem, we,e temporarily modified (i.e. accepted the CFTR gene), b<it tN' e ffects were
,eta1ively,hort-lived. Thisisbecauseour epith,elium cel!s arecon~11uallyreplaceda1 a steadyme,aodinCfpatientstheger,e1kallye11 gi"'-""edcel!s
arereplacedwithcell,withootUTR-codedp<imps. PatH'mswould,equi,eperiodkueatmentwiththe liposome---ONAcomplex ""'°""IOma i11tai11
theeflec! permanently
476
19.l Genetic techno~gy applied to medicine I
Social and ethical aspects of genetic engineering
and gene therapy
The benC!fits and hazards of gene technology
Genetically modified organisms have been produced for specific purposes, relatively quickly. where
8 Suggest what this proves possible. They offer enormous benefits However, geneticists 3re really producing new
advantage would rerult organisms when genes are transferred so there may be hazards as well as benefits.
fromthetramferof
genes for nitrogen
fixation from nodules Table 19.4 Thebenefitsandpotentialhaza,dsofgenetechoology
in leguminous plants to
cerealssuc:haswheat Hazardsthathavebeenanticipated
9 Occasionally • In a<ldiboo to human insulin production by modified • CoukJahafll\lessorganismsuchasthehumangut
newspaper articles Eroli,humangrowthhormooe,andbkxxlclotting <IMlerium f. colibe tm,sformed into a harmful
make criticisms of factor\llllhavebeenp,oducedbygenetka lly pathogenthatesc:"!)<"fromthelaboratOf}'intothe
genetic modification modified bacteria human population/
offoodorganfr,rm,
50metimes with • AnotherapplicalKlnofgeneticallyeng;nee,ing • Genesmovebetweenbacterialpopulationswith
alarmist headlines isbimemediation. Thisistheremmaloftoxk time.Coukltheantibiotic-resist.lncegenesin
Searc:hthewebfor compounds, carcinogens and pojlutanls, such a, pla,midvectorsbecomeacddentallytransfer,ed
artidesthatgive industria lsolvenls,cootamina tinggrnundwaters intoapatOO(jenkorganisml
ditferingvie1\!Sonthis andoudeo ilsp illsbygeneticallyengineered
Prepare the following bacteriawhichdegradethesesubstancesintos..fe, • Theaccidentaltransferofherbkide-resistancefmm
touseinadiscussion mok-cules.Theprocessi,,low. cropp!ants,, uchassugarbee~ toarelatedwikl
with your peers plantandthenontootherplantsisapossibility
a acorm1esummary • Suga,beetplantshitV<'beengerietkallymodifiedto "tikhcoold,esuM:in'superweeds'whosesprei>d
of the criticisms be'tolerant'ofglyphosateherbicideapplkations wouldbehardtoprevent
frequently made, Theseplantsareabk>toinactivatetheherbic;de
avoiding extremist whenitissprayed..,erthem,butthesurro,mding • Naturalinsecticidaltoxinengineeredintooop
language or weedscannotandarekilled.(Thisisdiscussed plants,whilstaneffectiveprotectioofmmbrowsir,g
further in this topic .) insects,mightham,pol li natingspeciessuchasbees
exaggeration andbutterlliestoo.
• Sim ilarly,natural,esistancetoattadbyche"ing
b alistofbalanc:ing • Thepresenceofthegene/0<1:hellttoxininthe
imecthasresultedfromtheadditionofthe
arguments in genesfo,Bttoxinintooopplants{potato, environmentmayleadtoinsectswith,esistance,
favour of genetic cotton,andmaize)bygeneticengineering.This
modification of food reducestheneedfore;rtemiveaerialspraying
ofexpensiveinsectkideswhkhareharmlulto • Thereatewellpu~icisedanx>etiesaboutfood
c arnndsestatement wildlife productsfromgenetka lly-eng ineeredspecies
of your own view Mightthefoodbecometoxicortrigge,some
on this issue • Thereareoo-goingattemptstousegene alk>rgic,eactioo,.Sofartherehasbeennoevidence
technologytotreatgeneticdiseases,,uchas of these
cystkfibros.is(page469)andseverecombined
immunodeficiency{SCID).ltishopedthatsome
cancersmayyieldtothisapproachinthe
""""
477
m Gene technology
Our laws are made by govemments and may or may not be ethically-based. -On some cases.
theymayeYenonlybeaviewfieklbyam;nority.)
Rel ig ion lnOU"sodetiesthen,arefola.,'l'l"Sofdiffell'l\l,eligionsardofnone.yetetl,ic,;applytousall
Cultu ral Many of these are littH, more than "fashooo,· that seemed acceptable at one time and whkh
maystill bel>eklur,uitka lly.
Whilst this may seek to give us an undmtanding of the oog;,,. of our worid and life. and how
thesewO<k.itdoesnotsuggesthowweshouklact
Our kel ing • Thesearelikelytobetheproductofou,earlyenvirooml'l\l,generalootlookand
o r conscience tempe,amen~ andoorind;vidualexperienc:es
On the other hand, the principles we do use in coming to ethical decisions are identified in
Table 19.6. You can apply them to the issues that gene technology raises, after you have read them.
ifyoucanacceptthemascomprehensive.
Regarding the c thks o f ge n e tech nology, perhaps there are two broad issues that societies
need to address in deciding what developments they wish to fund.
• Is there an important over-1"iding principle to be held to, that humans should not tamper with
nature in a deliberate way? Are the changes in biodiversity and genetic diversity that inadvenendy
flow from it completely beyond our current knowledge? Altemativdy, does all that we do
influenceandeventuallychangeourenvironmentsoourroleistoseethatouraaivitiesare
conduaedresJXXlsibly?
• To what =tent is gene technology a costly technology that is mostly beneficial to the health and
life =peaancy of wealthier people or perhaps to just those of developed nations? If the funds
used for gene technology were made available for the more basic problems of hollSing, health
andnutritioninlessdevelopedcountriesinstead,couldrnsdymorepcoplebenefitimmediatdy?
Alternatively, if we have the ability and resources to bring about beneficial development, is it our
duty to do so?
What does ) 'Our g roup think?
478
19.l Genetic techno~gy applied to medicine I
Tab le 19 .6Hhical,tandardstoapplyindecisionmaking
l lnfcnilil yandl\'F.
Fertilisation followed by implantation of an embryo does noc always proceffi in the normal way.
Either the male or the female or both may be infenile, due to a number of different causes
ln m.-tlcsthesemayindude:
• failure to achieve or ro maintain an erect penis
• structurally abnormal spenn, sperm of poor mobility. short-lived sperm or too few sperm
• a blocked spenn duct, which prevents semen containing sperm from being ejaculated.
479
m Gene technology
ln fc malesthesemayinclude:
e conditioruinthecervixUlatcausedcaUldsperm
e conditioruintheuterusUlatprevcntimplantationoftheblascocyst
• blocked or damaged m~ducts, which prevent Ule egg from reaching sperm
e eggsthatfailromarureorfailtobereleased.
In some cases. a couple"s infertility may be treated by the process of feniHsation of eggs outside
the body - in·l"ilro fcnilisa tion (lVF). lbe key seep in in- dlro fertilisation is Ule successful
removal of sufficient egg cells from Ule ovaries. To achieve Ulis, Ule normal menstrual cycle is
temporarily suspended witl, hormone-based drugs. Then the ovaries are induced to produce a
large number d eggs simultaneously, at a time controlled by the doctors. In Ulis way Ule correa
moment to collect the eggs can be known accurately. Eggs cells are then isolated from Ule
surrounding follicle cells and mixed wiUl sperm. If fertilisation occurs. the fertilised egg cells are
incubated so Ulat embryos at the eight-cell siage may be placed in Ule uterus. If one (or more)
imbcd Ulere, then a normal pregnancy may follow.
The firsi baby created in this way, called a "!est tube baby', was born in 1978. lbc process of;,..
cllrofcnilisationisilluscratal inFlgurel9.20.Today,thepr,:ccdure isregardcdasaroutincone
Figu,..,19 .20Theprocessof
'-'-""''""'""'.
hr
in·V!trofertoli<aboo bloc:ked.tempo,afi!y. (Pituit.uy
g!andaaivityisSUJ)!)(e<sedby
injectiooofaOO.mooethat bkx:ksthe 3Male(parmerordono<)
gooadouophin,eleasinghormooe) pmYicles"-""""sample
2S)<lthelicfSflisi~ed - ova,ies =:.::;::sedro
a,e,tiJOOlatedrodevelopmaoy
healthlestsp>rms
eggcelis(superovulation)
Genetic screening
mayalsobecanledootto
avoidanyct.ldrenwithserious
genelic:cood~ioo,
480
19.l Genetic techno~gy applied to medicine I
/u-vilrofertUisationraisesmanyis.suesforsociety. Jnfenilityisapersonalproblemtha!
generates stres.s and unhappiness in those affected. lnfenUe couples may seek as.sistance
and a cure. lbe medical treatments involved are expensive and their deployment inevitably
deflects finite resources away from other needs. Success rates are sometimes low. In the face
of the world"s booming human population, some argue against these sons of devdopment in
reproduction technology.
Another controversial is.sue is that current fenility treatments tend to create several embryos ,
all with the potential to become new people. HoWL·ver, the embryos that are not selected for
implantationultimatelymustbedestroyed - astressfultaskinitself.
ArgumentsforlVF ArgurnentsagainstlVF
• Forsomeotfierwisechildlesscouples.desired • Allowsinfertilityduetoinheriteddefectstobepassed
parenthoodm;,ybe;,d,H'ved oo (unwittingly) to the offspring. who may then
• Allows men and women ,u,m;ng cancer experienc:ethesamep<obleminadufthood
ITT'atmentsthepo,.sib ilityofhavingd,ildren • Excessembry05areprodocedtoensuresuccessandso
late,. usinggametesharvestedpriorto,adiatioo an embryologist has to select some new emb,yo(s) to
orchemotfie,apytreatments liveandtoallowthelate,destructionofothe,,potential
• Pennitsscreeningandselectiooofemb,yos human lives
beforeimp!antatiootoavoidaninherited • Mu~le p,egrlilncies have been a common outcome.
disease sometimesprodocingtri~ets.quadsorsextuplets.
• OffspringprodocedbylVFaremuchlonged-fo, lead ingtoincreasedrisktothemother"sfieatth.and
children who are more certa in to be loved and aisoincreasedrisksofprematureb;rthandbabH'swith
cerelxalpalsy.fo,ex.ample
• !Vfcanbeusedtotreatthemedicalproblem • H:canbearguedthatinfertilityisnotaM'aysstrictlya
olinfertility.Nobodyshouldbeallowedtodeny fieaH:h problem; rt may flave arisen in okle, parents WM
treatmentofaprnbk-mthatcanbecured choosetode!;,yhavingafamily(alifesiyH'iswe).
• Tfiereisanexcessolunwantedchildren.caredfo,in
orpflanagesOfinfoste,homes.Tfiesechiklrenmay
havebenefitedfromadoptionbycouples.childlessor
otherwise.keentobecaring'parents"
481
m Gene technology
~ - - - - - - - - - 2Heattreatmentof!hen)'Oflmemb<anet,;ndstheONAcopiesro~
fR
LlJ 3Selec!l'd.raooac!Nel;laWfedDNAprobes - - - - - - + 4NyloomembraneisnowOYerlaodv,i1h
~ r ~ ~ : = : i ~ ~ , ~ ~ , ,tONA ~~~ioo'"';,:isu:~'::;:=i.·
~
~
SX-rayfilmisdeveloped~ sOONinguptnepositioo,
ofthebar.ck(ffagments)rowhichprobesareattacl>ed
1====
Making radioai:t iv•ly labell• d DNA prob.. - -
• s;ng1e-manded ONA has tne at.lily to form a stable doobie strand with aoo1ne, ,w,gie strand of DNA. pruo,;ded
thebasesarecompk,memary(i.e.p,fr). llooe suandis"!abel~".thep«'Sl'nc:eolthepalrl'dmandsiseasilydell'Cted
• Short~!hsolsing1e-mar>dedDNAaremadein tnelabo<atoryf0<thispufl'O"-'.byenzymicalycombioing
aod!henaddi ngseleal'dnucleotidesooeat a time.inaprecisesequeoce
• Coosequently.!hebasesequroa'olpmbesisp<edeterminedaodknown
• All themdeotidesusedcootaonraooac!Nephosphorns(llP).acarbon(l"c)in thefiboseofthenudeicacid
backbonesothesubsequempositioosol1neprobes(andtheloca1ioool a complememarystraodolDNA,
e.g.ooan~memb<ane)canbelocatedbyau10<adio9rnphy.
A A G C G A h)drogenbondsbmkeo
/'
2Thetargetlor!heprobe - dooble-straoded 4RadioactiYepmbeisintmdlK!'d
ONAcootaoningthesequeocebeingstudied.
Figure19.21Stepsingenehcprnliling
482
19.l Genetic techno~gy applied to medicine I
The bulk of our DNA is not genes and does not code for pr(){eins. There are extensive "non-
gene" regions consistin g of shon sequences of bases repeated many times. Whilst some of
these sequences are scattered throughout the length of the DNA molecule, many are joined
together in major clusters. It istheselonglengthsofnon--ccxling, 'nonsense" DNA that are
used in genetic profiling. They may be referred to as sai cllitc DNA or as ,.,.rlabk number
tande ms repeat s (VNTR.;) . We inherit a distinctive combination of these apparemly non-
functional satellite DNA, half from our mother and half from our father. Consequemly. each of
us has a unique sequence of nucleotides in our satellite DNA (except for idemical twins, who
share the same pattern) .
To produce a genetic 'fingerprim", a sample of DNA is cut with a restriction enzyme which acts
dose to the satellite DNA regions. Electrophoresis is then used to separate pieces according to
length and size and theresuh isa pattern of bands.
To produce a DNA 'fingerprint' or profile·
I A sample of cells is obtained from blocxl. semen. hair root or Olher body tissues, 31ld the DNA
is extracted. (Where a tiny quantity of DNA is all that can be recovered. this is precisely o:ipied
using the polymerase chain reaction (see Figure 19.J on page 457) to obrnin sufficient DNA to
analyse.)
.l lbe DNA is cut into small, double-stranded fragments using a particular restriction enzyme
chooenbecauseit'cuts"closetobutnO{withinthesatelliteDNA.
J lbe resulting DNA fragments are of varying lengths, and are separated by gel electrophoresis
intobandswhichareinvisible,atthisstage.
-t The gel is treated to split DNA into single strands and then a o:ipy is transferred to a
membrane .
5 Selected. radioactively labelled DNA probes are added
to the membrane to bind to particular
12 Explainwhythe bands of DNA and then the excess is washed away. Alternatively, the probes may be labelled
rnmpositionofthe withafluorescentstainwhichshowsupunderultravioletlight.
DNA of identical 6 lbe membrane is now overlaid with X-ray film which becomes selectively 'fogged' by radioactive
twin5challenges an emissionfromtheretainedprobes.
underlyinga55umption .,. lbe X-ray film is developed and shows the positions of the bands (fragments) to which probes
of DNA fingerprinting, haveattached. lbe result is a profile with the appearance ofa barccxle.
butthatofnon-
Sieps 4 to 7 make up the technique known as Southern blotting and are Ulustratal in Figure 19.21.
identicaltwimdoes
483
m Gene technology
DNAprofilinginfo rensicinvestigation
rnmescl'n<' S1
1,themale(d')theparernofbothchik!ren?
Ex..minetheDNAprot;lesshown!Otheri,jlt
Lookat!hechildreJ1'sbaod,(C).
TheremaillOngbandsmatth!hoseofthebiologicallather
DNAfingerp,iminghasalsobeenw;cJe1yappied inbiology.lnomitholoq;,
torexample,DNAproli in gofr,e,tliogshasl'Stablishedadegreeof
'promisruity'inbreedingpai"s,themaleof'Mlichwas as.sunedmbe
!helatherof thewholebrood. lnb<rd,,!hepmductionofaclutthof
eggsisextendedowraperioo o l<iays,,,.,.;thc<>p<Jlat""1aJ1dfeniisation
:=(~t?~[.!,":,~:!::'Pf0¥ides theopporrunttylor
Figure19.22App1KationsofDt>1Aproliling
484
19.3 Genetically modified organisms in agriculture I
19.3 Genetically modified organisms
in agriculture
The ability to manipulate By the end of this section you should be able to:
genes has many potential
benefits in agriculture, a) explainth e,ignificanceofg=etic e ngineeri nginimprovingthe qualityandyi eldofcropplants
but the implications of and live,tock in solving the demand for food in the world, e.g. Bt maize, vitamin A e nhanced rice
releasing genetically (Golden rice™) and GM salmon
modified organisms b) O<Jtline the w ay in which the production of crop< wch u maize, cotton, tobacco and oil seed rape
{GMOs}intothe maybeincrea<edbyu,ingvarietie<thataregenetkallymodifiedforherbicide,e,i•tanceand
environment are subject
tomuchpublicdebatein c) discus, the eth ical and social implications of using genetically modified organism, (GMO,) in food
50fllecountries.
production
Bt maize
The soil bacterium &,cillus lhurin8ie11sis is a n3tural source of an insecticide known as Cry toxin
Genes for the production of this toxin have been transferred from this bacterium to seleaed crops,
=king them toxic to certain insects if the a nillllll should feed on the leaves, for =3mple. The
crops that have been genetically modified in this way are varieties d pornto, o:,rn and coaon. They
arethenreferredtoas Btcrops.
485
m Gene technology
Bt maize has been grown in the USA since 1995. The resulting crops have been used as biofuels
or in animal f=d only. Subsequently, Bt porntoes and Bt cotton crops have been approved for use
in USA . Bt cotton is the only genetically modified crop authorised for use in India and China. In
Europe, Bt maize is grown mainly in Spain, where the crop is used as animal feed .
TheadvanmgesforBtcropsdecectedtodateare:
• thelevelofroxin=pressionissufficienttobetoxictope.sts
• =pression of the Bt genes occurs within cells of the crops in such a way as to muse only in.seas
that feed on the plant body to be killed
• toxin expression has been modulated by adding tissue-specific promoters that have limited their
sitesofproduaion.
The use of these genetically nxxlified varieties appears to have been advantageous, at least during
the firs1 10 years. For example, the use of =ternally sprayed insecticides on corn and cotton crops
13 Explainwhyany hasfallensubstantiallyoverthisperiod.
threat to honeybees The disadvantage has been a more recent. increasing resistance to Be toxin among the insect
andbutterflieo;poses pestsofthesecrops.Asaresult.therehashadtobearerumrotheuseofexternally-applied
such a problem insecticidesromaintaincropproducrivity.
toagrirultureand Meanwhile, opponents of the use of Bt genetically modified crops continue to advance
horticulture arguments and anxieties about their adoption (fable 19.8).
Table 19.BAlgumentsad,ar,cedagainsttheuseolBtuopsandresponses
Argument Respon se
Po ll,,nfromBtcropscouldkillbutterflies.ir,cludingthe This ha,bpenqueried.sinc:ethepollenofBtplants
monarchbutterily. contaimaverylowleveloltmdn
Expefimentalevidencesuggeststhisisnolall'alistic:
danger. forthe,.monarchbutterfliesdonotconsume
maizepollenOfevenassoc iatewith theplants
BtgenesmaybetramferredfromBtmatzetonatural Alarge-sca\estudyinitiallylailedtofinde,,.;denc:eof
maize in neighbouring plots contamiJlillion
Subsequent experiments haw sh= some small scale
genetic:cootamination.Rl'S<'a,chcontinues
Btgenesincropsmaybelinkedtotheteeentdisease Thisisaiecentprob\emanditscause.whethe,dueto
of colony collar,se in honey bee h>VPS an unknown paras'te Of some tmCn. · unknown
Astudyby enagric:ultu,al,esearchgmuphasfound
noevidencethatBtuopsad,erselyaffectbees.solar.
c'°-o., j "• /,, "• leads ro night blindness. For children in many pans of the world
c~ aseveredeficiencyleadstototalblindness.Morereceruly,it
has been shown that vitamin A is crucial ro the functioning of
our immune system . Because of this, a deficiency d vitamin A
is the cause of very many childhood deaths from such common
[-~;~:~~c~o,j
We have already noted that rice is the staple for very many
people. Whilst the green foliaged rice contains beta-carotene,
the grain contains none. A genetically modified variety of rice
hasbeenproducedwhichstoressignilicam (andpotentially
life-saving)quantitiesofbeta-caroteneinthegrain.
Figure19.23Beta..:arotenea,asoo,ceofvitaminA
486
19.3 Genetically modified organisms in agriculture I
We earlier saw that herbicide resistaoce had hen, introduced into some crops using the
tumour-fanning bacterium, Agrobacterium (page 395). This soil-inhabiting bacterium invades
broad-leaved plants at the junction of stem and rool, forming a huge growth called a tumour or
crown gall. However, Ol:her pbnts, including the grasses are not attacked in this way. Consequently,
a different meihod of delivering the genes for storage of beta-carocene in rice grain had to be used.
Jn rice, the genes were delivered directly into the cells using =tremely small gold or tungsten
'bullets' that ,vere coated with the required DNA and then literally fired in with an approprfately
sized gun. This method of genetic modificatiOfl is used quite widely now.
A helping of about 300g of these genetically mcxlified rice grains, when cooked. contains
sufficient precursor of vitamin A to meec a person's daily requirements. However, there remaifls a
greatdealdoppositiontoit. Forexample,fieldtrialsofthiscrophavenOlevenbeenundertaken
in some pans of the world, including in countries where vitamin A deficiency causes hC3lth
problems. Opponents of the product argue that a more balanced diet that includes fresh green
vegetables is a healthier solution. One might query whether this response is rC3listic, given the
huge numbers d people that live in crowded urban environments where many items of a balanced
diet are nOl available to most people. There is no doubt that genetic nxxlification is a technology
with many opponents.
GM salmon
'Wild salmon (Salmo salar) have been genetically engineered to include growth-hormone regulating
genes from O,inook salmon of the Pacific ocC3n. This has resulted in a much faster growth rate,
and one that is maintained all the year round, rather than just in the spring and summer when
growth is fastest in wild and fish-farmed salmon. These GM fish grow to market size in 18 months,
ratherthanJyears.
Herbicide-resistant crops
Transgenic !lowering plants may be formed using tumour-forming Agrobacteri,m,. This soil-
inhabiting baaerium sometimes invades broad-lC3ved plants a! the junaion of stem and root,
forming a huge growth called a tumour or crown gall. The gene that induces tumour formation
occurs naturally in a plasmid in the bacterium. known as a Ti plasm id . Useful genes may be
added to the Ti plasmid (using restriction enzymes and Hgase - see page 4S8), and the recombinant
plasmid placed back into Agrobactcn"um. lben a host crop plant can be infected by the modified
baaerium. The gall tissue that results may be cultured into independent plants, all of which also
carry the useful gene. lbe plam wUJ then make the gene product. which wUJ be useful to humans.
Herbicide-resistantsugarbeetandothercropplantshavebeenproducedinthisway(Figure 19.24).
487
m Gene technology
Glyphosate is a powerful systemic herbicide. Once it has been absorbed by a green plant it is
tr:mslocated to all tissues and org:,ru and inhibits an enzyme essential for the production of amino
adds. lne outcome is that the whole plant dies. This enzyme is absent from animals. so glyphosate
has very low toxicity for animal life. On reaching the soil, glyphosphate is inactivated and rendered
hannles.sbysoil bacteria.
The gene coding for the enzyme that inactivates glyphosphate has been identified in these
soil bacteria, isolated and then transferred to the crop plants. These plants are now referred to
as 'herbicide-resistant" because when the herbicide is applied the GM crop plant degrades the
herbicide molecules and remains unhanned. Meanwhile, glyphosphate is also absorbed by we-eds
and kills even the largest weeds, including their extensive roo1 s,-·srems
- - glyphosateappliedinsolutKJO
viaa,p,ayboom
- -~ - - - J :1::.Kl:~~::ac1
withandistramportedall
aroundplants{systemic
flerbKide)
glypOOSateinhibitsenzymes
fmaminoaddfom,ati"" -
quic:klydisruptingtheweed
plant"smeta~ism
/,J/.~\'-11,,)"""L_- suga,beetgrowsfreeof
compelitiOllwithweedsfo,
lightandmineralOOns
Figure19.24Tn,atmentofaher00:Kle·resistants.ugarbestcrop
488
19.3 Genetically modified organisms in agriculture I
Table19 .99enefitsandpotentiald.rnge,so/growingglyphos.phate-re,istmtcrops
D• ngers
• lteradicatesweedsaroondsuga,beetcrnps,where • Fewerweedsperiesonfam>!andandkmge,pefiod,
k,,o;ofyieklduetocompetiboowithweedsis withoutweedsgro\\ingbreakswildlifefoodcha ins
veryh"]h • Selectiooofaglyphosate-resistantcmpp!antties
• Glyphosatehert><Ode istransportedallamundthe thegrowe,toooeparticularherb<odeproduct -
weedp!an~k illi ngeV<'nthelargPStweedswith chokeislost
extensive roots • W ill genetically mod ifted plant m..te,ial, when
• Glyphosateinhiiitsanenzymefottheprnductioo cOflSumedbyhumam,releasena.reltoxinsm
olessentialam;noac:ids inplants.Thisenzymeis othe<Wiseadvmelyaffectexistingenzymesystems
absentfromanimals,soglyphosatehasverylow inthehumandigestives)'Stem7
toxicity. • Genetic:allyenginee,edgenesmaybeYect0<edin
• Anave,ageoffivespra)'Sofherbicideareappliedto plasmodsrnnta ininganantibKltic....,,;,1anc:egeneto
suga r beetincoovenboMlcultivaboo,whe,easwith facilitatep,ocessesinthelab.Thislatte,genem>ght
GM beet a maximum of three spay; (and nom,ally beacddental!ytransfenedtohumanglilbacteria
oolyooe0<two)areneededbecauseofthegreater via food eaten
activityofglyphosate • 'Superweeds'maydevelopbyuoss-pollination
• Cerealsnom,a llyprecedespring-sownbeet.Cereal betweenherbicide-tolerantcropp!antsand
stubbleisanimportant'iOUrceoffoodforbird life compatibleweedspec:ies(fat-hen,seabeetand
inautumnand,,.inte,.Wrthglyphmatehe,bkode \\'l'edbeetarenatu,a llyocrnrringnearrelatives
app l>edtoGMbeetbeingsoeffective,the, eisno ofsugarbeet).Superweedswouklbedi/f,rnltto
needforweedrnntrolfr,stubbletheprec:eding er.tdic:atefromcmps
alilumn - leavingwildbirds'foodsoorcesavai!able • Thereisapossibilityolcros,-pollinafonbetween
longer. GMcrop,afldcon...,ntional andcrganic:cmps.The
• Glyphosateherb;codep<ovidesgood"'l'edcontrol, maintenancecfsuffi6entdistanc:eorthedevis.ing
alk,\\;ngrnftivatk,mbydiscsottines(mme,va!ion cfeffectivebarrierstopre...,ntorreducepollen
til!age)rather thanbypk,oghing . Theses)'Stems tramfersbetweencmpsm..ybedifficulttoachie""
f,a...,fewer harmfuleffectsoosoilo,ganisms, • tfglyphosateheibic:idedropletsdoreachhedgefCM'
inueasemoisture,improvescilstructure,and p!ants(orfurther)theirslzeandchemic:alactivityin
reducetillagec05ts plantsmeamtheyaremore likelytododamage
• Glyphosateherb;codeisappliedas!argedmplets
fromrnarsenozzU.S,rather thanasafinemist
oltill)'dropletsthatarepmnetodriftonto
surroundinghab;tat,
489
• Gene technology includes genetk engineering, DNA foodplants\\ith herbiclde resistance. Insect resist ance or
seque ncing and genetic fingerprinting. Gene technology has i~foodvaluehavebeenproduced
~iCillionsinbiotechnob9',:medicineandthepharmaceuticals • DNAsequenc ing irwoM!sthecreationofgenomiclibraries
industries,agricult1xeandforensicsdence. ~tedviologf of the precise seque nce of nucleotides in samples of
generates many benefits for humans. but there are potential DNAofindiYidualorganisms. Thenucleotidesequencein
hazards,too,sogenetechnologyraisiesethicalissues. the 1\tiole human genome was the product of the Human
• Gen etk e ngineering involves the transfer of genes from Genome Proj ect. Many other genomes have been completely
onespeciestoaoother,possiblyunre!atedorganism.Genes sequenced.too.
aretransferredbyinsertingDNAintoavector,typicallya • Gen etic profiling involves the analysis of DNA to identify
plasmid , a tiny ri!)Q of dooble-s1randed DNA obtained from the individual from which the ONA was taken . It is used to
abacterium.Thegeneand thellectorilfecutbymeansofthe e;tablishidentityandthegeoeticrelatednessofindiYiduals,
same rest riction e nzyme. forming compatible sticky ends at for eMJTiple in forensic science. The ONA samples are cut
the cuts. The gene and plasmid are then brought together and by restrittion e nzym esandthelragmentsseparated
joinedusingtheenzyme llgase.Analternativevectoristhe Electrophoresls isaprocessusedtoseparatemoleculessuch
nucle icacidofavirus. as proteins and nucleic acid fragments (of either ONA or RNA)
• Bacteriathathavebeengeneticallyengineeredtocarry a ontheba!.isoftheirnetchargeandsize
newgeneare ld e ntlfl edbytheuseofplasmidsthatcarry • Genetic disord ers are heritableconditionsthatarecaused
genes for resistance to two antibiotics, known as R-plasmid s bya~ficdefectinageneorgenes. Mostarisefroma
Altematively,af lu orescentmarker isused mutationinvolvinga slng le gene.Gene ticdisordersaffect
• Foragenetobee,;pressed(transcribedintornessengerRNA} about 1- 2 percentolthehuman population and include
apromot ernormallyneedstobeattachedandactivated sickle cell anaemia, haemophiliaandcystlcflbrosls.lt is
One eT.ample is the lac promoter mech.:nism present in certain po5Siblethatthe symptomsofageneticdison:lerlikecystic
bacteria. fibrosis may eventuaHy be ove«:ome by gene therapy. The
• A gene may be constructed from the messenger RNA Human Genome Project has made it possible to screen for
that it codes for, using the enzyme reverse transcriptase. thepreseoceofthemutantcysticfibrosisallele.Consequently,
Alternatively, copies of genes may be built up from affectedpeopleandtheirrelativesmayberelerredto a
nucleotidesinthecorrectsequence. Thisisworkedoutfrom genetic counsellor whose role is to take a detailed case
theaminoa.cid se quenceoftheproteinthegenecodesfor history and to provide information from which lt-.e. clients can
usi!)Qthe ge ne ticcode. make choices.
• Crop irr1)1"overnents are also attempted Uf genetic
• Ge netkally modified ba.cteri a are now used to make engineering. Projects have included herbicide-resistant o~
v.aluable products such as human insulin (for the treatment
seedrapeandsugarbeet,insect,resistantmaireandcotton
of diabetics), human growth hormone and several other
andvitaminAenhancedrice. Theresultshaveoftenbeen
hormonesandenzymesofuseinrnedicine, agriculrureand
c.autiously and eYen aitically received. sometimes with possible
otheriooustries. good rei!SOflS. Geneticengineeri!)Qremainsatechnoloqy
• Gen etkally modified eukaryotes ilfe harder to produce. withmarl)'opponents.very fewcounuieshaveapproved
Theycarrytwocopiesofagene, unlikeprokaryot es, so the genetically modified oops for convnercial production
engineeri!)Q processes are often more difficult. However,
490
Examinationstyleqoostions I
Examination style questions
1 a) Ahusbandandwife1it.oalreadyhaveachildwithc.ystic c) Improvements were found in the vision of all the
fibros.is{CF}electedtohavethefrsecondchildtestedfor participants, but the small number in the trials made most
theconditionwhilestillafetusinveryearfypregnancy. oftheseimprovernentsnotstatistic.allysignificant
The results of the test, a DNA banding pattern, were Suggestwhythesetrialsweredesignedtoincludesuch
discussedwithageneticcounsellor. a small number of participants [2]
TherelevantDNAbandingpattemproducedby (Total: 7)
----,--
electrophoresi1i11howninFig.1.1 (Cambridge International AS and A Level Biology 9700,
Paper41 Q5November2011)
first
d,ild
3 a) Four naturally occurring enzymes make possible the
geneticengineer\activities.Listtheseenzymes,their
1l"'"""'fil~of
mOYementof
fragments
sources, andthereactiontheycatalyse
b) lnthein-vitrosynthesisofhumaninsulin,acopyofthe
[8]
"' Fig.1.1
i) Making the insulin gene from mRNA compared
toobtaininggenesdirectlyfromrelevant
chromosomes [3]
Withrefereoc:etoFig.1.1,explainwhy,
ii)Thechoiceofinsulinobtainedbygeneticengineering
ii) thefetuswilldevelopCF [1]
forthetreatmentofdiabetics,comparedtotheuseof
ii) thepositionsofthebandsofDNAofthefir~
insulin from animal sources [3]
childandofthefetusindicatethatthemutantallele
(Total: 20)
for CF has a deletion in comparison with the normal
allele [2]
b) ExplainbrieflytheneedtodiKusstheresultofthetest
4 a) Give an illustrated arrount of the separation of DNA
fragments by electrophoresis. [6]
withageneticcoumellor. [4]
b) Explain·
[Total: 7]
(Cambridge lntemationa/ AS il!ld A Level Biology 9700, i) Thenatureandsignific.anceof'variablenumber
Paper41 Q6November2009) tandem repeat' sections of chromosomes in ONA
fingerprinting
2 In humans, thegeneRPf65encodes a prote1n responsible ii) ThestepsintheproductionofaDNAprofile [10]
forregeneratingvisualpigmentinrodandconecellsafter c) OutlinethediverseapplicationsofONAprofilingin
theyhavebeenexposedtolight.Arecessivealleleofthis science and society. [4]
genecausesimpairedvisionfrombirth, progressing to (Total: 20)
complete blindness in early adulthood. This condition is called
LCA 5 a)Outlinethestepstocropimprovernentbygenetic
ln2008,trialswerec.arriedoutintothepossibilityand modific.ationwit hrefereoc:etoc.anolasugarbeetcrops
safetyoftreatinglCAusinggenetherapy. and herbicide resistance
a) Suggest and explain why LCA is suitable for treatment b) Identify both the potential benefits of genetic engineering
using gene therapy. [3] offoodcropsforagrirulturalproductionandthepossible
b) Sixadultswiththisconditionwereusedinthestudy. hazardsthatmayarise, illustratingyouranswerwithrefer-
Genetic.allymodifiedadenoviruses(atypeofvirusthat eoc:etocurrentexamples.
cancauserespiratoryinfections)wereuseclasvectors
Thevector1wereinjectedbeneaththeretinaofone
eyeofeac:hoftheparticipants.
Suggesttwoway;inwhichthegenomeofthe
adenovirusesusedasvectorswoulddifferfromthatof
normaladeooviruses. [2]
49f
Answers
1 Cell structure
I Trarufcrd~(respir:,tiof\l.fredingornutrilloo,mctabolism,
excrcdon,moverncni andlo.ornooon, re5PJmivencsoorsensi---
1Mcy, rcproductm, growth and devclopmcnl
2 a) l mm•IOOOµm
·-·
SincelOOO+ l OO· IO. l Ocdbol l OOµmwiUJi. r along
"-"'""*"...)
-·.:<··--
""'loclu, onddvomatw,
(_,...,...,.._
<hloropl••
·-thll-11<.
Plant cells Anlm•lc.ils
Cellwilll Cellulose cell wall Nocelluk>secellwall<
~:,:::,g, ...
Manycelficontain
chloropl;r;ts,siteol
photo,;ynlhe,is
Nochlo,opl•sts:
a.nimalcellsc;orn>t
photosynthesise ~--
...r<h .............
~«n..,....,_..,_
~.,.-n>ilo<hor,:lrio,-
492
Answers
e.g. c.ilcium c.irbon:i te and magnesium carbon:ue, in the 2 a) Acaulysii.oasub11t:once tlu1 a!1ersthera te ofachemlcaJ
Mlells of non-vcnebr:ue animals. mostly marine but some of reaction. bu1 remains unchanged at the end.
fresh water and terresuial habit:iu, and aa ch:iJ.k and Jimc..-
.,1ene nxks. which are formed from foei!i,cd shclli.
Typiulyamet~,suchas
2 Sec &ic~nd chemiwy for biologists on the ID. pL,tinum,inllfinely-dMded
3 SecDi.'laccharides.page}S.
4 Sec Polysaccharides,1»3cs36-9,pank\J larlyF',gurc2.9 _,.
s1a1 ...... g.uplitinisedminer;,I
5 Sec The function:d group, of sugars, page 32. Abletowithsundh igh Prote111ist.rsilydenaured
6 A polymer L!; a large organic molecule nude of repea ting t,,mper.itur.s, highpressures byhighte<nper.llurl'.for
subunits known as monomers, chemically combined
togcthcr. Among the carbohydnucs. starch, glycogen and
;,nd .. xtremesofpH,if
=""'·
cellulose arc all poly men buill from a huge number of E~melysi,K•llc-most
molecules of glucose (the monomer, combined together in a,esi,Killc \oonetypeof
different ways in the three. disdnci: ivc polymers) subs\utemolecule
7Secpage32andFigurc 2. 18,page45 ',) ,nrtoalme
8 Given the n~turally occurring pool of 20 differcru type'! of
amino add5. In a polypepdde of only S amino acid rcoidues
there can be 20'diffcren1 typcsofpolyp<..-ptide, or 3200000.
In a polypcpi:ide of 25 amino acid rc•iduc• there can be 2(!S
different type•.
In a polypcpi:ide of SO amlno acid rcoiducs there can be 2~
different type•. A polypeptide o( more than SO amino acid
residue•i•.byconvention. callcdaprotcin
So.thcrc<!;virtuallya!mostunlimitedstnxturalvariation
po61libleintheprotcinsofccl!s. Rcmember. aa thescqucnce
ofaminoacidrcsiducschangcs.socloco thestructureand
propcnieaoftheresultingl)l'OO!in.
9 Sec page3SandFigu re2.19.pagc46.
10 Sec Figure2.22,page47.
11 1he term macro,nolcci;le means 'giant molecule'. We have b) 96/40•2.4crn's-•
.1Cfflthat polysaccharidc,(e.g.ccllulosc)andpra:cins(e.g. 4 Pre--incub:1.1ion is required to ensure that when the reaci:ants
l)'!IOZ}m,c)arcmadcu p ofv.utnumbcn:ofrepcating smaller att" mixed the re,mioo occurs at the known, pre-selected
493
Answers
r:I free water molccules is low) will show a g.ain r;1,., ter t.uicset.
molcculcsattheexper,seoithec:liluteglucosc lol1.11ion. b) 0 For example, inthela)..,,-r:lccllsformingsperm,in
the!este.s
ll) For example. in the bone marrow where red blood
cdls are formed or in thc generm:ive layer of the skin
atthe bascoftheepidermis
494
Answers
b)
TheprodllCti 2idenliulcells,
effliwiththe
di!)loidch==
numbe,.
Producesidenbcal
4non-identialcell:s,Nchwith
thehaploidchromo50ffll!rumber.
Proouceshaploidcells.Contribvt"'
1 11
.... __.. ,...................
. ,-..""v""'"-...-*
4&-.;,Fig urc 6. ll.pagel20.
SSeeFigurc2.19.page46.
__
... . ,
di!)loidcells togenetkva,iabHityby
P«mitsyrowth • redocingchromosomenumbe< 6 RNAlnvolv.dlntHnoaiption
within multicellular byl.alf,permittingfertilisation, mel5efl9e<RNA Seelhe~ofprotein"Yf'thm.Sugeone
--
aganisms,.ind andthecombiNtionofgene, (mRNA) tril~tion.p•ge120
asew.ol~roduction fromtwol),llrents; RNAlnvolv.dlntr.n.i.tion
• pem,ittng r a n d o m -
of l),lltemaland matem;il
me55enget RN.A mRNA is ro.v a linur molecule in tht cytoplnm
(mRNA} K>ngwhichribosornesrn<l'>'e,·!Nding'the
• recombinatiooofsegmentsol
g,entticcodeardlm1'cri>ingthtinforrniltlOll
indMduolimatefNlindp,itemal
Into• ~ne.tr""luence of alTWIO ;icid residues
(,,primarystructureofth tproteio}
homologol,schromosom"'
duringcrassingover. lransfe,RNA Seelhesl"'Jl"ofproteinsynthtsis.Stagetwo
(tRNA.) aminoKmactivation.page121
S Sec Table S.2 Common carcinogens known to increaac muta-
(lon ratcs and tM likelihood of rnn«'r, page lOS. 7 See alsoFigurc6.IO. page 119andFigurc 6.t6. page 124.
6 46><2•92 a) Gly(glyclne),Asn(asparagine),Pro(proline),
7 :i) See 1able under pan (b) Phc (phenylabnlne), V..I (vallne), Thr (lhreonine),
Your pie' chan ~houkl compri,e 9eCIOnl "'ith the His(histidine),Cya(cy.sieine)
follO'•dng angles at the centre: Prophue 2S2". Meo.phase b) CCT, TIA, GGA. MA. CAA, TGA, GTA, ACA
36°.Aruipha5cl8", T ~ S4o. c) SeeTriple1:codes.axlonsandanricodons.page 122.
495
Answers
.~ T5 of a sunflower stem (low power)- plan diag ram I O Cultivation and cropping of plants to supply focxl for !he market
placeinvolvesthe"harvesting'dorganicmaner(andthenutri-
entionsbuiltintoi!duringgrowth).lfthesupplyofessential
mineralsthatplantsrequireisnotmaintainedinthesoil,the
'"~~-rn
~a/trace producrivityofthe land wm rapidly fall. Consequently, Jargely
solublenutrientsareaddedasanificialfertilisers spread on the
..
soil as granules at the timed pe-:,k crop groMh. Alternatively,
vascular farmyard manure is applied to the soil at the timed ploughing.
This decays slowly and releases nutrients more steadily.
"""' I I Mitochondria produce ATP (the energy currency molecule)
ATP transfers energytoenabletheactivetransportofsolutes
Iscale:10mm
-.,-' ""
cambium
in the sieve tubes. The presence of mitochondria o:infi.rms
thatphloemtransponisanactiveprocess,u.singenergy
transferred during respiration.
12 a) lbe functioning o f living ceUs (e.g. in the phloem tissue)
wouldbeimpairedasproteinsweredenaturedandcellsur-
face membranes disrupred Translocation would stop. 'W-ltet
4 Your answer should refer to the features of their Slructure
flow in the xylem vessels would be unaffected.
(see The movement of water through the plant . pages 135--8,
b) The concentration of sugar might be lower in a sieve
panirularlyFigure7.8),theirintimatecontactwiththesoil rubenearthebaseofthestem,ifstarchisbeingstoredin
solution and to their position on the growing roots. lbe thecellsatthat poim.
signifieanceofagmdieruinwaterpcxentialbetweensoil l .l a) Sugardeliveredbythephloemfrcmtheleavesisstored
solution and the root cortex is expbined in Upmke of as starch(insoluble)onarrival. l>fany oftheions absorbed
water - therolesoftheroothairs(pagel36). fromthesoilareusedinthemetabolismdtherootceUsor
5 The casparian strip blocks the apopla.st pathway of water arecarriedtothestemandleavesinthexylem.
movement across the cortex at the endodermis. b) Sugarsproducedinthelightinthechloroplasts
6 As temperature i = = , evaporation of water from the accumulareinthecellsoftheleafbeforebeing
translocatedawayto"sink'sites.
surfaces of the cells in the leafi==· Al the same time, the
==tofmol.ecubofwaterisincreased(t:heirkinetic
energyi.srai.sed),sodiffusionoccursmorequickly.Also,as 8 Transport in mammals
temperaturei==,thehurnklityoftheairisdecreased I For =ample. it means that blood cannot be directed to a
becausewarmairholdsmon,w:itervapour.Sothegradiernin respiratorysurfaceimmediatelybeforeoraftersen>icing
watervapourbetweentheinterioroftheleafandairout.sideis tissuesthataremetabolicallyactive(andsohaveahighrate
increasedanddiffusionofwatervapourisenhanced. of respiration). Rather, the blood circulates randomly around
7 SeeThetranspirationstream,pages 140--1. the blood spaces and blood vessels.
8 Transpimtionisadirectconsequence ofplamsa-ucture,plant 2 Sx 106 cellspermm3 isequivalentto5>< 109 cellsperlitre.
nutritionandthemechanismofgasexchangeinleaves.ln So, inSlitres(i.e.thewholebody),thereareSx(Sx to9:> -
effect,thelivinggreenplantisa"wick'thatsteadilydriesthe 2.S>< 1010cells.
soilaroundit.Putlikethis,transpirationisanunfonunate If a red blood cell lasts for 120 days, divide the total number
o:insequenceofplantstructureandmetabolism,ratherthan of cells by 120 to find how many. on average, need to be
avaluableprocess.ltmeansthatao:irutant,adequatesupply replaced each day:
ofwateriscriticaltoplantgrov,th: ifthewatersupplyfails, 2.5>< 10 10 + 120 - 2 x 1,Jlapproximarely
plants cannOI: move elsewhere, as animals tend to do. Pulmonary circul ation Systemiccin:u lation
9 a) Plant growth is dependent on a supply of chemically
Carriesdeoxygenatedbloodunder Carriesbloodooderhigh
combined nitrogen (for protein synthesis) of which
highpressurefmmtherightside pressurefromtheleftsideofthe
nitratesaretypicallythemostreadilyavailable.However,
ofthehearttothelung, hearttotheall thebodyorgaos
nitrates are also taken up by microorganisms a nd may
be released in the soil at times cxher that when plant Retumsoxygenatedbloodunde, Retumsdeoxygenatedblood
demand is at its peak. They are also very soluble and lowe,pressuretotheleftsideof unde,lowerpressuretothe
are easily leached away into ground water in heavy rain. the heart rightside oftheheart
Bytakingupnitrateswhenevertheyareavailable(and Coosistsofpulmonarya,teries Consistsofaortaandarter"-'S
storing them in cells). plantgrowthcanbemaintained deliYelingbloodtolung,.cap;ll ary deliver;r,gbloodtothebody
b) Plant rootsaremetabolicallyveryactiveandrequire netwmksseMngtheai,sacsof orgaos.capillaryr,etwo,k,;
oxygen for aerobic respiration and ATP formation (for the lungs. and pulmor,ary ve;m seMng all the tiss\ll'S and cells
ion uptake, for example). Waterlogged soil Jacks soil air carryingbloodb.ad:totheleft oltheorgansofthebody.and
and the essential gas, oxygen side of the heart venacavacarry;ogbloodback
totherightsideoftheheart
496
Answers
4 ..) lhcvdodry:ondpr=,urc<Xthebkxxla.1itCl'IICflltheaort:a 7 Tuexnon-daakstrand<kttptheheanvalvcflapspointing
arcbodthign,bu:asitisabouttore-cntcrthehcanthe inthediredionclthebkxxlflow.They..wpthevalvcsturnin8
pre55Urc lsv.,ry low. Ho,,,._....~,do8e 1othe hea/1, lhe vdoc- inside<>u1whenthe~risesabrupdywithinthe,'mlride;,
i1y ofthe bkxxl in the vciru is •imilar io1h:u ofthe main ar- 8 a) The pressure in the aorta is alwaf!i significantly higher
1~leavi",1:thehean.Thisisncc,,ssary1ocruurcadequate than that In the a tria lxcause blood i, pumped under
Vt.'l'IOusrt.•(urntobalanr<."therardiacoutpot high prCMureintothe aorta and, during diastole and
b) 11-.e blood speeds up on leaving the caplllarie•because atrial systole, the semilunar valves prevent backflow from
thetotalc!'OU-sectionalareaofthevenu lesismuch leM the aort:i. Meanwhile, blood emers the atria under low
than thecapillaries. lhetotalcross-sectional area of the prcs..,urcfromthevciruand thepumpingactionofthe
venous sy,i:em further decreases as venulcs join up to atria is slight rompared to that a the ventricle. which
form the veirn. gcneri,tesour'pul.,e'.
c) The velocity a the bkxxl is at its lc"":est in the capillaries. b) Pressure falls abruptly in the atrium ~ ventricular
Capillaries are the suwles1 blood vessels bu1 their tOOll ,y,i:ole is underway a.1 :mial cliaMole begins then.
~are:i.isl)-l'ically800timesgre:i.t~than c) 1be.K'lllilunarv.ilveintheaortaonlyopenswhenthe
thatofthcaorule:aving the hem(diametef"ofabout ~ure in the ventride,r e:,:c=d,, th:1.1 of the pres5Ure in
25mm). The slow bkxxl flow in !he caplllaries alLOW5 for
effidente:,:change ""'·=
d ) Attheendofsystole,prcssurcinlhevemridcsishigh.
5 a) The components of the blood that are nol found in ti...ue When ventricular diastole rommencc5 the bicuspid v.1ke
fluid arc the blood prolein,, mainly a lbumin, and red will only open when pressure in the, ventricles fal[g
blood cell• and platelets. (White blood cdb arc found in below that in the atria.
1he(issuefluid - seepagel5J). c) About 50 per cent of the cardiac cycle is given O\'Cf
mdiastolc-the resting phase in each heanbeat. The
heart beats throughout life and takcs limited rest at !hC$C
~ ~ ~ . /'.. /
,
.. / ' ... }-
~~~)\
I
497
Answers
4 About 10-15%. Land-living organisms obtain oxygen from the The fate of the control group is followed in comparable
air,butdiffusionwithinanorganismoccursinsolution,so detailtothatofthetreatedpatientsinordertoevaluatethe
thefirststepinoxygenuptakeinthelungsisthedissolving efficacyofthe=perimentaltreaunent.NO!ethattheuseof
ofoxygeninthesurfacefilmdwaterattherespimtory such a control group means that a treatment is withheld from
surface. As a consequence, the respiratory surface is kept people who might have benefited. This raises obvious ethical
moist and water vapour will evaporate as gaseous exchange
occurs.SeeFigure9.6,page176. l l • Life expectancy has greatlyincreasa:1 - inthepast, people
5 Gasexchangeistheexchangeofrespimtorygases(oxygen
and carbon dioxide) between cells of an organism and the • The development of antibiorics has led to a decrease in
environment:cellularrespimtionisthecellularprocessby deaths from infectious diseases
which sugars and other substances are broken down to • Improvements in knowledge and training of medics means
releasechemicalenergyforothercellularprocesses.Gas that symp1oms are more correctly attributed to actual
exchangeisaconsequencedcellularrespimtion.
• Dietarychanges
6 Sec Movement by bulk transpon page 93
Peoplehadlesssedentarylifestyles
, ~~~,'--,
,ffoct
~ ,-, o~
d ,~oo-,.~,=
~M- ,-, ~ ~ ~ ~ 12 • Deliverscarcinogenstotheairwaysandairsacs
Su,fa.c:eareaof Ahugesmfaceareaforgaseousexchange • Delivers carbon monoxide to red blood cells where it
alveoli (Approximately50m 2,abouttheareaofa combine-sirreversiblywithhaemoglobin,preventing
doo~estenoiscourtl) oxygen transport
• Raises blood pressure
Verythin,llattened(squamous)epithelium{Sµm
• Triggers va.soconstriction cl blood vessels
thicklsothediffusioopathwayisshort
• Stimulatesthesecretionofviso:iusmucusbygobletcellsof
Capilla,ysupply Anetw00:Colcapilla,iesa10<1ndeac:halveolu, airways
toalvl'<>li rnppliedwithdeoxygenatedbloodlrom • Inhibits the beating movements cl cilia in the epithelium
thepolmonarya,teryandd,a;nir,ginlothe of the airways
pulmooary ""'"' majnta;n, the cement, atk,n • Leadstoblockingofairwaysandtheaccumulationofdust
gradientsoloxygenandcarbondKJxide and carcinogens in bronchioles
Oxygendissolvesinthewate,l;nir,gthealveoli • Causes loss ofnaruralelasticityofsmallestbronchioles and
aOOdiflu=intothebloodinsolutioo
E!astic:fib,esin Stn>tcheddufinginspj,ation,theythenrecoil, • Jnflamesthebronchiandcau.sespersisterudestructive
coughing up of phlegm
walls lac:ilitatir,gexpiration
• Leadstonarrowingofairwaysandeventualdestructioncl
8 Carbond,oxide,sanac,d,cgaswh,ch.,fnwereto airsacs
accumulate in the blood. would alter the pH of the plasma • Leads to the pos.sible formation of a rumour - Jung cancer
solution. The normal pH of the blood is 7.4. For life to be
maintained it must remain within the range pH 7.0-7.8. This
is largely because blood pH affects the balance of essential 10 Infectious disease
ions which are transported in the plasma solution. Efficient I Present in the wall of the stomach are millions of tiny pits
removalofrespiratorycarbondioxidefromthelungsisas called gastric glands. The cells in the gastric glands make and
importanttolifeasefficientuprnkeofoxygen secretethecontentsofthegastricjuice.About6oocm3 of
9 ln1950theincidenceoflungcancerinmenwashigherthan gastricjuiceissecretedpermcal,secretionbeingcoordinated
in women , in keq,ing with the fact that the percentage of the with the presence d food. One of the components of gastric
adult female population that smoked cigarettes was lo wer, juiceishydrochloricacid(strength0.15M)ataboutpH I.
about 40 per cent compared with 6o per cent cl males. This is sufficient to create an acid environmeru of pH 1.5--2.0
lntheperiod194S...-1 975thepercentageofmalecigarette for the contents of the bolus, the optimum pH for the protein
smokers declined slowly whilst the percentage of female digesting enzymes of the gastric juice. 1be hydrochloric acid
alsokillsmanybacteria,amongorherusefuleffects.
cigarette smokers increased slowly. Associated with these
2 In cholera, chloride ions move from epithelium cells into
changingpattemsincigareaeuse,theincidenceoflung
the lumen of the gut by active transport caused by prO!ein
cancer in males staned to decline after 1985. whereas in
pumps in the cell membrane, driven by ATP. Water moves
females the incidence of lung cancer was slowly increasing
throughouttheperiodthisstatisticwasrecorded. byosmosis(aspecialcaseofdiffusion),inwhichmetabolic
cnergyisJX>tinvolved.
10 Acontrolgroupinamedicalinvestigation:
comprises the same number of people as a patient group } The insecticide DDT is harmless to humans at concentrations
selectedforan=perimentaltreaunent thataretoxictomosquitoes,butappliedatthese
is as comparable with the patient group as possible, as concentrations it rapidly collects in body fat and is
regards age, gender, health, occupation and general life transferred from prey to predator through the food chain,
experiences of its members steadilyincreasinginconcentration. ltaccumulatesatthetop
consistsofpeoplehavingthe'condition'forwhich of food chains at concentrations that may cause harm. The
treatmentisbeinggiventothe=perimentalgroup.
498
Answers
stability of the DDT molecule means that it only very slowly by the tobacco industry for an =cep1:ionally long time. Now
biodegradcs. This persistence is why it is so effective, but is we know the various reasons why cigarerte smoke triggers
also why it is a health threat. By means of her book Silent malfunctioning of body systems, ill health and diseases of
Spring. it was Rachael Carson who publicised this problem. varioussorts.So,statisticalconfidenceinthepossibiliryofa
4 These figures show us that, wha!ever the level of TB causallinkisaspringboardtofunherinvestigation,notproof
infection in a region, about 10 per cem of those infected die of a relationship.
from the condition. Also, almOllt twice as many people in 8 The full range of reasons are contained in the text, How
Africa are infected with TB as in Asia and the proportion af eradication came about, on pages 210. Individually they can
the population of Europe with TB is far lower than in either be shown not to apply to one or more of the other diseases
oftheothertworegions. discussed in this topic.
ThesedifferencesinprevalencearemOlltlikelytobe 9a) Expo.,ureofpathogenicbacteriatosub-lethaldosesof
accoumedforbythreemajorfactors: antibiotic may increase the chances of resistance devd-
Housing,whichisrelatedto agenerallevelofpoverty oping inthatpopulationofpathogens.
and unemployment. Where the prevalence of TB is high b) Byvaryingtheantibioticsusedthereisincreased
theenvironmentalandsocialconditions(panirularly likelihood of killing all the pathogens in a population
housing)arelikelytofavourtransmissionofthe including any now resistant to the previous antibiotics
bacterium. Countries with high unemployment and low used. This approach works umil multiple-resistance
wagesandcountriesexperiencinglocalorcivilwarsare
strainshaveevolved,suchasinstrainsofaosrridium
more likely to have a major pan of the population living
difficile and Staphylococcus a!lreus, for example.
incrowdedandunheahhyconditions.
10 Antibioticsareaddedtothefeedofintensively-rearedlive-
Access to medical care. lbe levels of diagnosis and
treatment of illness vary substantially. More of the stocksuchaschickensandpigs. Heretheyreduceorprevent
populations of African countries are without access ro theincidenceofmanydiseases, andtheanimalsarefound
effectivehealthinfrastructuresthanintheothertwo to grow better with them. Possible dangers arise from the
regions, for example. Where there are fewer doctors over-exposure of bacteria to antibiotics see Figure 10.29
or care centres perheadofthepopulationorinrural (page21S).
communities that are widely scattered, the chances of
diagnosis,treatmentandpreventativemeasuresare 11 Immunity
minimal.
lbe cost of health provision. Some countries cannot I SeeWhatrecognitionof'selrentails, page220and
affordrobuymedicinesorvaccines,oronlythe Figure11.4,page221.
relativelywell-offcanaffordthem.fowcountrieshave 2 Antigens may be present more or less anywhere in the body
the equivalent of a health service so the poor do not get that can be contaminated from outside the body. Antibodies
exist in the blood and lymph and may be carried in the
5 HIV/A!Ds cause:s pank,.1lar problems for the people and the blood plasma anywhere that blood 'leaks out' to, including
economies of less-developed countries such as Zimbabv,e or sites of invasions. They also occur on B-lymphocytes.
Zambia because it almOllt exdusively handicaps the young }SeeFigure8.2,pagelS}.
adult population at their time of greatest economic activity 4 Spleencellsandrumourcellsculruredseparatelyandthen
and family building. mixedtogether.Mixtureisculruredtoproduced'hybridoma'
6 The pathogen is able to survive within only one specific hos! cells. Resulting cells are separated and tested for antibody
andthereisnoreservoirofthepathogeninod>erorganisms required. Positivehybridcmacellsareculruredfurtherand
(forexample.anyofthesurroundingwildlife). By contrast produceamibodyinsignificantquantities
the TB bacUJus =ists as a reservoir in wild manunals (such 5 a) The =istence of memory cells avoid the steps to the
asbadgersintheUK) production of activated T-lymphocytes. The particular
7 When two events (A and B) regularly occur together, it memorycell,oncere-activatedbythere-invading
may appear to us that event A causes event B. This is antigen, switchestoproductionofan=cessofthe
not necessarily the case, there may be a common event appropriate plasma cells and helper T-lymphocytes.
b) The initial response to an amigen by the inunune
thatcause:sboth, forexample,oritmayanentirelyfalse
system is slow. For sufficient antibodies to overcome an
correlation. An =ample of the latter are those infants (very
infection takes weeks rather than days. When memory
few) who develop the symptoms of autism shortly after cells are present, many stages of the initial response are
the normal time in childhood when the M.'dR inoculations omitted and antibodies are quickly assembled.
is administered. It was scme time before detailed studies 6 lmmuniryistheresistancetotheonsetofdiseaseafter
established that MMR inoculation and the onset of autism infection by harmful microorganisms or internal parasites.
were not causally linked. Long-livedspecificimmunityisaresult oftheactionofthe
Becausecorrelationdoesnotprovecausewasjustonerea- immune system and may be acquired naturally by previous
son why Richard Doll's amassing of statistical evidence of a infection,butcanalsobeinducedbyvaccination. SeeA
link between smoking and iJJ health was successfully resisted summary of the types of immunity, page 2}1.
499
Answers
Mltibodies,eceivedvia
........
monoclotllllantibodie-;
7 a) ln ll:5pnlion. all the hyaogen alOm.1 arc gr;,duaUy fCITlO\'ro
from glueo5C, catlly5cd by dchy<rogenax enzymes. The
h}'Q'08Cflaloms2"':odckd!0hydrogenaccep(On,WWlly
lmm ,..... lty theplac:enta NAD (nicotlnamlde adenine d inudaitidc. page 23S),
forming reduced NAD. NAD i, a coenzyme 1hat wofks
Immunity with specific dchydrogenasc enzyme,, in the oxidation of
sub6tra1c molecules by the removal of hydrogen.
h) Dcc:arboxylationistheremoval ofcarbonfromorganic
12 Energy and respiration
compounds by the formation of carbon dioxide. For
example.gluCO!ICcoruisuofsixcaJbonatoms.All
six carbon a\Onl$ are removed at differcm M.agcs of
ro:spir~tion. ooc :u a lime. and given off a.s carbon
dioxxie. A specific decarb=ylax- nlZ)TilC i5 involved ln
c:och case. The 6m: decarboxybtion ln aerobic rcspir:uion
2 The illlpOflant fe:aturn of ATP ~ !Nt it is:
ocrursinthcrcactionlinkingglycolysiswiththcKrebs
cycle, when pyn,wtr i,, convened IO a "'-"a-carbon
• a aubslan«" dt:ll moves ~ily wilhin cells and organisms -
molecule . The other decarboxylalion reactions of aerobic
byfacilit:ueddiffusion
respir:llionoccurinsiepsintheKrebscyclc.
• formed in cdlular respiration and takes pan in many 8 a) A subinratc is a molecule that i5 thc starting point for a
reactions of metabolism biochemical reaction; it forms a complex wi1h a specific
• ablelotTansferenergyinrelativclysmallamount.,, cnzymc. An intcrmediatc is a m o lecule formed as a
suffidenttodriveindividualreactions. component of a metabolic pathway.
J Sec Extcn$ion: The r~piration of fats and proteins, !)"ge 239, h) Sec Glycolysi.5. pagc 243, and The link reaction and the
andExcrction,bloodwaterbalanccaodmctabolic""'5te, Ktcbscyclc.pages24}-5.
pwticularlyFigur-, 14. 14.p:ige?:97. c) SccExtcnskln: Respir:uiona.sascricsofrcdoxrcactioN.
4 C,5I IJ1COO H • C,(il-!3,0, page 238.
C,6' 1.w):+ 2302 -4 16CO,+ 16H,O + Energy 9 a ) Glycogen
RQ•:::::
b) Starch
10 ~onlcrcactiorurcquire=ergyinput,bcca usethc
produc.u have more poteruia.l cnergy man !he reactants - !ICC
RQ • l 6+2J•0.696 FigurcJ.l.page57. Note: 1healtemative1ypcofrcactionls
5 ;,) onc in which the products have ]es., potemial energy than
thcrcactan1s.11tcsercactionsuan sfercncrgyas hcatand
work andarccallcdexergonic reactions.
11 Glyrolysis(thcconvcrsionofglucosctopyruvate)which
occuf"!linthccytosol(theaqueouspartofthccytoplasmthat
surround., organdies such as the mitochondti.t) docs no(
rcquireoxygcn rotcornpktion.
12 U..C of a single wa.ter bcuh :u a favour:i.bk tcmper:uurc -
pcrhapis ~ oe; rcpc!ition of the expcrimml vdth a range of
dilutcglUC06CK>lutions(thcsub:5u:r.1c,).
13 l.aru,te and a fflla.U amount of ATP""' the fiNI products of
an.acrobicrcspir:Uioninmusclefibres.
14 In the a ~ of oxygen rcduccd NAO accumula1cs and
oxidisedNADrescrvcsareusedup. Inthc ab.!cnceof
oxidisedNAD,pyruv:n eproducrionbyglycolysi.sslows and
.stops. .so subscqucnt stcp, in respiration must stop too.
time ahergemiinitionldays 15 Presence of aercnchyma tissue of .stcm. lcafand root: ability of
b) s«dlingAh:asapatterndRQch:ingelnCllrly roots1orespircan.acrobica!ly - withrclativelyhightoleraocc-
S<"Tmination typia,l of sccd.s with a food store ofcthanal.
«lnSUting m3inJy d carbohydrate. (Actually. tin., cbta 16 lnthcrcspiromctcr.thcfarsideofthcU-1ubc1Nnomctcris
isfrorng,:nnin:i.tinawheatfruitswhkhoontain7ffll> 1hecon1rolmbc(A). Hcre,conditionsareidcnticalto!hose
carbohydr.m,,2'*'lipid(oil)and 12'*.pro!Cin).
in thc rcspiromctcr tubc, but in thc former. no living material
SecdlingBhasapattemciRQch;,ngeincarlygcrmi=tion
i5presen1. Hov,cvcr,anychangeincxtcm:al1empcraturcor
1yp;c;,J ofSttcls with a f<xxl .w.., consisting mainly dlipid.
pressurciscquaUyexpcricncedbyboth 1ubes,andthciref-
(Aaua!ly, dtis cbta is from germinating !bx 5'eCd.1 wluch
contain 2'K carbohydrnre. 55% lipid (oil) and 20% pmtein) fecta on 1he level d m.anome1:rk fluid arc equal and opp<>
6 Thcfollowingareproduceddw-ingglycolysl!:,rcduccdNAD sitc.andtheycancclom.
(c), ATP(c)and pyruvate(g).
500
Answers
501
Answers
--
4 When a diabel:ic patient aucsscs their blood glucooe by myeli11sheath imy,elil1shtath ~inated
means of a bioscnsor, 1hc measurement is d the a mount of 2
glucose prc-sem at 1he moment the reading is taken. When
readingsaretakenofgluoosclevelsintheurine,thereha.,
Stimulusdet.ct.d ] Reu pto,
l la<atlonof
Mechanoreceptor1
bee,, an inevitable lag time before blood glue°"" level is
rcflcctcdinthe urine.Golorimetricreadingsinvolvealevelof
Stretch receptors, I Sl<eletalm,,sde
subjcaive judgment that the bioscnsor a=ids, too.
5 u) 1b: force that drlves ultrafiltration in the glomerulus
posilioo
I e.9.musclespindles,
propr10Ceptors
is th<- blood pressure generated by the muscles of the Bloodpressuie I Saroreceptors
ven1rides. Thispressureishcightencdbytheefferem
am,riolc of the glomerulus being of a smaller diameter Thermon,cepton
than the afferent arteriole.
b) Water, useful ioru, glucoec and amino aci<b, along with lnternaltffllpe<ature l~,~=us
6 a) Brush border is made up of mkrovilli. Microvilli provide
Che..-eaoptor1
a v.ully increucd Mlrface area ol cell •urfacc membrane BlcodO,,CD_i,H' I Ca,ot,dbody l ca,otid-..y
in comact " 'ith the fil!r.ue. In the membrane, there arc
proccin pumps which actively and selectively absorb
usefulmeiabolitcs.
b ) Sccl'acilitatcddlffusion.pages8l-2,and l'igurc 14.19,
p;ige3()1. J lnpositi,,-..:f~theeffcctof adcviationfromthe
norrnalor5elconditioniatocrcatea tcndcncytorcinfc,rc.,
7 lna cou111cr-cun'efllflowsystcmflui<bflowinoppo0<ite
thedeviation.PoliitivcfcedbQckintcnsificsthcoon-ective
direcrions in parallel and adjacent tubes. So the limbs d the
actiontakenbya control systcm,lcadingtoa"Vidousciocle'
loopofHenleandthcv:asarcctailll ilocc:ur,ibcside,ithc
siruation. Imagine a car in ,1:hich the drlVC'f', 5e2t """" stt on
loopofHenlcarcbo<hoounter-currcnrsysicrru.Exdtange rollers(ratherthanbcingsccurcdtotheOoor)bcingdrivcn
o.eur,ibe!ween1hesesysiemsandagr:1.dlenli,,m:1.i111ained a t speed. The •lightest applica tion of the fOOI brake cause,i
along the entire exchange Jllffacc. the driver to alide and to press harder on the brake a• the car
8 a) A negative feedback sy~cm consist< of a receptor, a st:uu to slow with an cxtJcmc ootcomc.
coordinator and an effcctor. ln osmorcgulation we have: Biologicalexamplcsofpo15itlvefccdbackarcrare,but
Re<:cptor:osmorcccp(onin thehypotl,alamus one can be identified at the synap,c. When a wave
Co-ordinator:hypothalamu5andpo11tcriorpituitarygland ofdepolarisation(anervclmpulsc) takescffoctinthe
from where ADH i.s released post.synaptic membmnc, the cn1ry d sodium ion• trigger• the
Effcctor:collcctingduct5 of the ncphron•inthemcdulla
entrydfunhcrsodiumions a1agrcaterrate.n1isisacase
ofthekidnt.-ys.
b ) 1bc osmorccep(cn ln the hypod,abmu•. together with of po11itive feedOOck. The clcpolarlscd stale i• established and
the rec,,pror•in thea01taandcarotidancrie,idetcctthe theimpubemovcsalongthepos tsynapti<:nt(."mbrane.
change in the water content of the blood. Nen,-., impulses lnneg:itivefccdbacktheeffectofadevia tionfrornthe
from the receptor• are transmitted to thc hypc,thalamu• normal or S<."I condition i., to create a tendency to eliminate
andpiruitary gland. thcdeviation.Negativcfeedbacki•a !X'rtda!m().S{a!l
9 SccFigure4.17aodFigure4.18(pages88and89). con1rol sysrems in living things. The effect of negative
10 Stomarn, open and doee in re5pon,c to ch:rnge11 in rurgor fcedb;icki.storcduccfunhcrCOJTcctiveactiondthccon1rol
in the guard cells. Sec 1llC opening and c!O<.ing d stomata, systernoncctheset-pointvaluci.srcachcd.
p;tg""30S-9andAbl!cisicacidandwaterwessintheleaf 4 :a) Sec,1h, reMingpotential. page 316.
b) S=1'heactionpotential.pageJ17.
"""""· S Sec,The refm,ory period. page 318.
502
Answers
6 Sec Speed of conduaion of the action pocmtial. p;,ge 319. 2 Anuj,orcvmtofint"'Jlh;,seisthereplicationofthe
7 a) Tr:uwnincr ""D'll:Ir><:el llff" pnn,ccd i-1 the Golgi :ipp:u:,rus du-omosomcs. By the omission of this ~lllge ~'CCn mciosu
in thesyn:,plicknobandheklintinyvesldespriortouse. l andmeiolisll.thech--numberishalvcdinthe
b)Sccl'igure!S. IO, page 32 1 fourcellsproducedfollowingmeioticccl!di\'Uion.
M Scc Tiieulm,.sm,crureofskelernJ muscle. and Figurc, IS.II. ., a) Mctapha se landll
15.12, and Fig IS.IJ pages 322--324 b) Midproph:iscr
9 lhecffectsdhormone-sarerestrictedtoceUswithspecificrecep- c) Lateprophasel
ton on the ccll !iUrfuce membrane to which they attach or with d) Anaphasell
whichlheyre:i.ct.ootheyaffectonlythctaigetcell.!ortiw.>es. e) Anaph.asel
IO ln ncgatlvefecdhack1heeffectofadeviationfromthe 0 Telopha se ll
oormal Of set condition is to cn,ate a 1endency 10 elimirune
Mitosis - •
the deviation. Negatfrefeedoockisapanofalmo,n:i.11 r.pliutM d ivision
control 5Ystenu in Uving ~ - lbe effect of ~live Consequences Allcel.carrythe • Ensuresmaintel'IM".:e
fe«backistoreducefunherC01Te<:tivcactionof1hecontrol Hmegenetic ofthechtomasome
system or,ce the set-point v:i.lue is ~heel. inlormaticna.the number in body cells
lnposltivefo:dxtcktheeffeadade-viationfromthenormalor existingcelhlrom lromger,eratiooto
setrondltion is to create a tendencytorclnforoethcdeviation wh«:h they are gene,aticn
l'o5itive feedback intensifies the <:errcctivc action tiken by a !OfJlled,~r>dwhich • lsaswrceolvariitioo
controlsystcm,lcadingtoa 'vidou,drcle'situation. Biological ttley,harewith (importantlorttle
cxamplesdpo,,itivefe«lbackarerare,butooecanbeidentified ,umiurnlfr,gcells 1,1.JWalofthespecies
at the syn::,p,<e. When a waved depolari.'Qtion (a rio:,rve impulse) inachangir,gwo,ld)
takes effect in the postsynaptic membrane, the entry of oodiwn Twoidentic.alcelh. fou,.non-itlen~ c.Hs.
ionstriggenthcentryof further «xllumlonsatagreate,-rate NCh ... rththe eachwnhthehaploid
II a) Al about the mkl-point in the mcnstnial cycle (<by 14) d,ploidnumb.,of nutmefofchromosomes
thehighandri,singln-doloestrosmsuddcnlylilimulates <h-e
thei«retionof lrl and,toaslightlylc:ssa'"extmt. Significance • Penni1'gr<M'th Contri:>utestogenetic
FSH. by the piruiwy gland. u-i stimulates ovulation ardrepairwithin variability by·
(the shedding of the mature se«>ndal)' oocyte from the
Graafianfollide)andthe.sccondal)'oocyteisrelea.,ed
muJticdular
organism, ,,_
• rfllucingthe
..~tionof
~gmf!ltsofincfrvidial
16 Inherited change matema,l;ind
I a) Scc Chromosomes occur in pair$, page 96. pate.-nilhomologous
b) MitOl!is andmciosisaredivisionsclthcnucleusby vcry chiolTl050rneSdu,ing
precise pro.es.ses; they ensure the correct distribution of
chromosornesbetweenthedaughiercclls
Titccl:tught<.""rcdlsproduccdbymitosis haveasetof S Thei<leathatlhecharacreristicsclpareni..wcrcblmdedin
chromo5omes identical to e ach O(hcr and to the parent theiroffsprlngwasdisprovedbyinvestigatingthcprogenyof
cell from which they were formed . ln growth and crosscsoforganismswithcontrastingcharacterisdcs(suchas
"tall'and'dwarf'peaplant:'l)througha~andsometime,,
development it is es.sential tha1 a!! cells carry 1he same
lnfonrunionutheexiatingcelli,.SimUarlywhenrepair =bscquent gener.,tions. l=ge samples were used and the
of <bmaged or worn out cells OCC\11"$, the new cdb &ff" experirnmt,wererepeated,confirmingth:1.tthcobscrved
r:itio<werestatisticallysignificant:.
exact~ofwhattheyreplace
Mcio<!i5 occurs in the life cydc of all orgmi!:ms that 6 ~ nn expect a r:itio of 3, I among the progeny only if
reproduccsexually. lnmriollisroord:tughtcrccllsare thr~ conditiomarcmet;
produced. each with one member d each homologous • f'"'1ilisation is random - each fenilis:i.tion jg lndcpendcni of
pairofthech=~escitheparcntcell,knownasthe all the others
haploid (,i) ~tatc Halving of the chromosorne number is • thcrearecqualopporrunltiesforsurvivalamongd1colfsprng
csscntialsince atfertilisationthenumberisdoublcd. • large numbers of offspring are produced
503
Answers
In breeding =perimem.s with pl3nts such as the pea plant, • This also means that. by elimination. Mr and Mrs Lee must
exactrntio,maynotbe obrninedbecauseofparasited3mage, be the parents d the blood group B child.
due to the acrion of browsing predators on the anthers or • Mr and Mrs Santiago are blood groups AB and 0. which
ovaries in some flowers or because some pollen types fail to migluproduce:
betransportedbypollinatinginsectsa.,succes.sfuUyasothers. AB x 00---> blood groups A or B
7 ThelayoutofyourmonohybridcrosswiUbeasinFigure1 6.14 Since Mr and Mrs Lee are the parents d the blood group
(page355) ,buttheparentalgeneration(P)wUlhavegenotype,, B. Mr and Mrs Gerber must be the parents of the blood
(ifyouhave chosenCtorq:,=tthegeneforcoatcolour): group A child.
cRc'xc"' cw • Byelimination.MrandMrsJonesmustbetheparentsof
whereCRrepresentsthealleleforredcoatandC"'represents the blood group AB child.
the allele for white c001. The gametes the parenml generation 9 3) SeeRed-greencolourblindness, page359.
produce will be: b) See Sex linkage, page3S8. and Figure 16. 17,page359.
l O The recombinants among the F, progeny are those with
The offspring (F 1) will have the genotype c' cw roundseed.sandgreencotyledonsandthosewithwrinkled
andthephenotypewillbe"roan'. seedsandyelloweo1yledons.
lnasiblingcrossoftheF,generationthegametescibolh
siblingswUlbe:
~ c' { c""'
504
Answers
12 a) BBSS,BB&!:,Bb.SSorBbSs 15 Firlilcalculaiethc1J.,utistic.
t,) Fromatestcross,usingaspar*lthat huarcdsponed
co.u.Amongthcpuppie,iprodu=:l,lhcpre,en«ofrcd
coa1 orof•pottedpanernindicales !hat thcaherf"lre!U
~-----y.,...,-,.,..,.~~~~~~=~~
::- ~o =i
spaniel l< heterozygous for that cluraeler. tall. a><YI
I :i A mrnant organi,m (or cell) is one carrying ahered genetic
material which makes it different from its parent (or from its tall, termiri.al 1
precursor cell).
14 a) dwarf,~ial 1
dwari,
Soi•0.384andthcrcare~~offrttdom.
From a table of the dlstnbution of 11 (se,, page 36-0. the
probability(P)ofobl:aininga deviation a.s large a.s(orlarger
than)theonewehavebychancealonefallsbet"-·ecn1he
probabilitylevclsclP•0.90 andP •0.95.
This is not significant - there is no depanure of the observed
from thee,ipccted values.
16 Gern: mut:Uioru arc due to ch:mg"" in thc ~mce d ba5C$
in the ONAola gene. ruchas when one t:we i., replaced by
another in the coding Slr:lnd of DNA th.:11 makes up the line:u
sequence of bases.
17a)Llzand Diana
b) O David and Anne
JO James, Wdlbm, Arthur and Fredrick
c) 8 (Richard and Judith, Anne, Otarles, Sophie, Oiris,
Sarah,andGail)
d) James and William, Arthur and Diana, etc.
18 A dominant allele is expressed in all individuab !ha\ inherit it
(evenifonlyhe1crozygousforthat allele),whereasarcccs-
siveconditionappcarsonlyinoffspringthatarehomoeygous
for 1ha1 allele (and therefore with parems who were both at
Jea51carr\er1oftheallek ~areb1ivelyrareevent).
19 The genescoritrolllngrhe production of the blood pro1cin,
concernedinhacmophilbareloc:i.tedOlltheXchromoeorne
Haemophililt is caused by a recnsive allele. As a rc,uh,
haemophilia 15 brgelyaconditionofthc, male since in him
a•ingleXchrortl0$0II\C'carrying~defectiveallele(XhY)
will rcsul! in dlsease. For a female to have the haemophilia,
she must be homo~ygous for the recessive gene (X~ xl'), a
condition 1h:it ls frequently fatal in u/cro.
505
Answers
506
Answers
c) 1he annc,wion.5 to the curve arc an opportunity to • Dd"orc,tation-Dcstructionof trec,idesuoys the habit:1.lll
speculate on: of nu merous &pccics d aninuls, p!ant.1, fungi and t»ctcria
• onthe initi.o.lrapidriscinnumbett (continuingirn,- whkhthetcforcdeclinc innumber.i.
migntion«, initial abscncc of rompclillon for abundant • Dcseniftca1lon-Thisisspttded upby:
reK1Urceoi, lack of presencc of predaton when popula - • overgrazing cl land.
tio n ncw/ low in number;,) • deforc,tation.or
• on sublicquent stabilisation of population numbcrs • climate c hange.
(balaocc betweenreproductioo/ pred:ttionprcs,ures, • Exccssive application of pesticides in modem industrial
theideacic:arryingcapadtyfotthathabitat). agriculture. Pcso:icidcs have improved productivity Jn
6 1be Simpson Divcrsity Jndcx for 1hi, habit:lt: agricul1ure.buttheir u.., hasttduccdbiodivcrsity.
• Air pol lution - The lcvcl.s of carbon dioxide in the
Sped1t1(nooflnclividuals) nl,, - 1)
atmo,pherc have been rising since UX" lnduwial
Revolution in the developed co,mtrio of the world.
1ltls gl05 is a major contributor to global " " rming and
--~
ddUUC1ivcdimatcchangc.
lotal{N) • Water pollut lon
'·' - - -~
In (n -l}• 1980+ 156o + 90•36JO.
D• 9 5 x ~ - ~ - 2.46 -•+--chromosome-,
ing.ometo· in~-podoong
7 The original data in Figure 18.11 were oblained by a team p,O<b:ir,g<oll ,..,
ofpre-Universitystudentsin:mactivitytheydesignedand
carried out (independently of teachcrs) as the culmination
of a field oounie. Rcspond to this quescion in ~ke-nun...,...
Selccttwosnu Utcarruofso:udem,;,cachtotadtlcthe
y
calct.tl.ation of onc of the di,.-=iity indices. Then. by
groupdiscu91ion,=:11.luatcthecffCC1ofage/stagcofdune
development ondiver.;ity. PrC'le(ll your data and conclusions
as a ~er or display ncwspapcr iu,m
K Precisely ddi.ned and internationally agreed scientific names
i,-rJ----
facilitate cooperation between ob,;c,rvera by ide ntifying the
CJCact species that is being inveso:igated andrcponedon.
9 TI1e central rol umn needs to oi:an with 'Domain: one of the
three major forms of life". The left· and right-hand columns
need toparallelthisadditionwith 'Domain: Eukaryota" .
10 Sedimcnt:uy rock layers are usually laid down under
water. often under a naerobic condi1ions. and formcd
from sediments washed in from the land. They arc addcd
10. }'Carafterye:ar. Objects that falltothebottom(for
example. dead organisms) a rc covered. compresocd a nd
eventually their molcculC!l react with or arc replaced by
miner~! ion,. Foss ils laid down in the deeper s tra ta of
sedlmc ntaryrock.,areolderthanthe fonilsof thehigher
(m o rcreccn1),1rata.
Dead organism that fall dsewhere are much more likely
to be dismembered by scavengers and/or decayed by
mkroorgani= beforeanyfossilisationispo,siblc
l! Thechiefecosya:tern.sare:
• Tundra. e.g. alpine tundra. which occurs on the highest
mountai,u. well a bove the 1rec-linc
• Grassland. e.g. so:eppc, pr.i.iric and pampas. or sa.w.nnah,
whichllltropic2.lg=,.sbnd
507
Answers
....... ,. ......... ! .!I Thc stocks in seed banks mus< be '1able. 0v"1 very long
T,,,...,1,1,nd,quwti,c c..,.,iv.b...ilng
prog,,m,,..sofz-...d pc,riocls,viabilityofs=d.sdecrea.scs,.sofreshformedsceds
•.db.nk11tbo\linic.l arcne«lcxlregularly.Abo,geneticviabilityismaintainedby
g,rd..,, thisproceS11becausenewseedsaretheproductsof sexual
Mabitatsthatm,alreadyrare OtigOllcllly.zooswe,ecollectioo, reproduakm and 1"0 maintain the diversity of the gene poc,l.
areespeci.illyvulr1"rableto of largely\lllfamiliaranimals
r>aturildisaster-meli.,bitats keptforcu~withSttle
themseM!Saieusilyloo.t.if conce,nforanystressc.o......d. 19 Gene technology
--6
arangeofe,camplesarenot butnowaptivebreed,ng I Viral DNAtakcsov"1thcribosomalm:1.chineryofthc
preserwdasnaturere,erves programmesmakegoodll'il'of hoslccllcausing1heproductionofvir:a.lpro11cinsd1iuarc
enzymes. Viral enzymes aiusc the replic:Hion of viral DNA
Whenah~td~ C~breediogmiin~i"" and the production of other rnoleruln that make up new
the"''holeccmmunity;.lo<t, lhegenet,cstockolrare•nd viruses~Figurcl0.19.page206).
threateningtoi.-.::rusethetotal end..,geredspecies 2 a) See Nucleic acids- 1he informa1ion molecules,
,,...mbe,olen(bnge,edspecies f>'l8"5IIO-l3,andFigure6.J,pagell3
h) See Table l9. l,pagc4SS
Arefugefo,endangered Thege,,eticproblems ari,ing
3 a) Astickyendconsistsofa short"""!uence ofexpo5Cd
wildl ifeal!owstlwsespeciesto fmmindMdualzoosh aving
ba,iesofunpairednudeo1:idesfonningasingle-strancled
leadnatural l,Ves in • fam iliar v,,rylimiteodnumberstoKta,
~tension at the end of a length of DNA. h is created
environme,itfor"hid,tfleyare parentsisO'o'ercomebyinl<'f·
by the action of a panicular restriction en;r;yme Iha!
1daptedand tobt!ap•rtof iooco-<:>peution(-ifldartifi.d.al
r«0gnlscsa specificbaseoequcnccin alength ol DNA
theirnonn.ilfooddwin< inwminationinsomecasf'>l
and cuts the DNA in thi5 Wtinctive way. See abo
Thewildlifeofareoervenwy Aninwlsinzoostmdto Figurcl9.4.pagc4S8.
be monitored for Nriyw•rning havesignrfantlylongerMe h) Sticky ends attach 10 ..,.rn omer by complemcn1ary
ofanylurtherdflerioration .:irpecundesandireiYail•ble base pairing. lhey are held together by the formation
innumbersof.th .... tm.d !Dpartic~teinbreed,ng of hydrogen bends between complementary bue.s. The
spKiessothatremedial!ileps p,ogrilfflmesfofmuchlonger enzyme lig:lsc then analyses 1he formation of C-0 bond,,
a,nbetiken thanwildanim;,lsdo betwecnthesug:,r-pho,iphatebadcbonesolthetwo
Theolfsi:iringolordangered ( aptivebreedingproblems,fo, ONAwancb.
spKiesarenurtu,.d inthei, mostspeciesitis apploedto, 4 a) See Table l6.2,pagc3S7.
natu,alenvironmentand gain have been highly successful, h) See Table l9.l,pagc4SS.
all thee~rienct'Sthi, nom,a lly althoughthe)IO<lngdonotgmw c) The bact"1lal chromosome is a single circular molecule
brings,irduding the.tequi,ition l.()inthe'wild', so then!isles, ofDNAwithinthccytoplasm,butattachedtothcccll
ofst:ilsfrompa,..ntsandpee!'S opportunitytoobse,wandlearn surface mcmbnne at one poinL Plasmid< arc tiny circular
11,oondthem lmm~entsiOdpeers mole,culcs ol DNA, fr=-Hoating in the cytopla11m of aomc
Thereisanesubli5hedtradition 8re,edingprogrammesgt!ll'f.ate bae1erla. Theyrepliaitcindepcndcntlyofthcbae1cril,l
olmJ1nuiningreserw,;and he111thyindMdo.iilsingood <hromooomo.
protectedareasinv•riousparts nurrber,lo,attempts.atre- S The genes of prokaryotcs :ue generally <:29"1 to modify dun
oftheworld,sott-.ismuch introduct,onofen<linge,red tho&cofcukaryoiesbecausc:
e,cperiencetoshareonh1M1 to <pKieston;,turalhibit•ts.• • prok:uyO(CShaveasin31c,cirrul:irchromc,.,omc,1t0only
mi,nagethemsuccesrlully particularlychalleogingproc""' one copy of a gcneha,,to beengi~ in!Othcir DNA,
giventhatnatur1lpftdatcrs whercascukaryotes havetv,oalldcsofcverygenc
abour>dintheselocation,
508
Answers
• plasmids, the moSI useful vehicle for moving genes, occur 9 Many newspaper repons are often rather superficial. You
inprokaryotesbutmostlydono(ineukaryo«,s - the mayhavetoamplifythecriticismsslightlyinorderrobeable
excqxion.s are some fungi, including yeast, and a few plants to introduce balancing arguments. Your own opinion needs
• tr:mscriprion of DNA into RNA in prokaryotes does not stating simply, with one or two dearly explained reasons.
require the removal of 'non-infomt3tive DNA' (introns), but 10 The presence of one or more sons with haemophilia among
in eukaryotes it does the progeny of the members of any generation in a family
• bacterial walls can be crossed by plasmids (after suitable treeisthewaythatca1Tierscanbeidentifiedamongthe
treatments). parents
6 A mutant allele is one of a pair of alleles in which II Thisisahighlypersonalissuewhichneedshandlingwith
(exceptionally)therehasbeenachangeinthescrucmreof great sensitivity. This is certainlythecaseforanycounselling
the DNA. Typically this involves a single base substitution, a team whenever this situation arises. They will need to en.sure
translocation, a transposition or an insertion or deletion of a thecouples'preferencesemergeandtakeprecedencein
baseorshortsequencesofbases ensu'gd'scuss'ons.
7 Apersonwithasinglealleleforcysticfibrosisisa 'carrier' A!Jyoutacklethisquestioningroupdiscussionwithpeers,
and does not =press the disease; whereas someone with a it is best if each individual =plains their own view and is
single allele for Huntington's disorder will become affected listened to carefully. It may be that a consensus concerning
bythediseaseevenrually. asuitablere.sponsecanbea1Tivedat - butyoumayfind
8 The fruits of the cereals (which include wheat and rice) thatyoucannotallagr=. Jnthissiruation ,youshouldtryto
are the staples of human diets all over the world. They understand the view of those you do not agr= with, rather
provide the bulk of essential energy- richfocxls. lne ability thangetinroconflictwiththem.
of leguminous plants to fix atmospheric nitrogen for amino 12 DNA fingerprinting relies on the fact that each individual has
acid and protein production enable these plants to grow well unique DNA. The exception is identical twins; their DNA is
without the addition of(expensj,,,e) nitrogen-basedfenilisers. identical. The DNA of non-identical twins is not.
halsomakesfocxlproductsfromthem(peasandbeans, I} Flowering plants are a major component of human diets and
forexample)relmj,,·elyrichinproteins. Cereals that are very many are insect-pollinated. Honey bees and bunerllies
geneticallymodifiedsothattheycouldalsofixnitrogen(if are some of these insect pollinators. A reduction in their
they were created) would grow well without nitrogen-based numbers would lead to fewer seeds being fonned. The
fenilisers. They may also add even more to human nutrition supplyoffruitsandseedsforcultivationandasfocxlwould
by helping to the overcome protein deficiency as well as be threatened.
meetingrnlorien=ds
509
Index
dihybridCl'OiSS 36()-{,3 ascendinglimb ( loopofHenle) 3()2
abioticenvirorunent 384 dominam 352,357,369,370.4@ asexualreproduction 103,344
AB0bloodgroupsystem 3So6 monoh}1>ridcros, 35()-...8 aSllistedreproduction 447
absciskadd 309. 336 multiple 355
absorptioospectrum 165 receSllive 352. 357,369----70,4@ All's,,eadenosinetriphosphate
accidents 194 allopatrks~ciation 408 All'- ADPcycle 236, 237
ocetykholim." (ACh) 319 alphacells 293 All'synthase(All'ase) 246,247
acetylcoenzymeA 243,244 a -glucose 31 atria 164,165
acidophil"s 431 a -helix 47 atrialdiastole 166, 167
octiofilameots 249,324,325-7 altirude 162- 3 atrialsystole 166.167,168
action poteotial 317---9,320 alveoli 174.178-80 atrioventrirularnode(AVnode) 168
actionspectrn 265 American grey squirrel 448 atrioventrirularvalves 164----6
octivationenergy <;8---9 aminoacid.s 44,45,~,51,llS-19,HS Au.straliangrasslands 384-5
activation step 222.226 autism 209
activeimmuoity 199,231 allllllOflia 297 autoimmune deficiency syndrome
activesite 59--{il.94 amniocentesis 471,472 (AIDS) 204--7
activetr:mspon 78,90-2,128,30L316 Amoeba 2.10. 89,434 autotrophknutrition 234,262. 431
adenine 110, 111,112.113,118 amylase 61,62,63-'l autotrophs 'Z72
adenosinedeaminase(AD) 469 amylopecrin 38 auxin 336.338
ad.,nosioetriphosphate (ATP) 21- 2, amylose 38 axons 92,312,313,314,316,317- 18
111- 12,235,236-7,238
ATP- ADPcyclc 2.J,6. 237 anabolicreactioru S6,57,235
ATP syntha"" (ATPa"") 246, 247 anae robicre-spirntion 25}-7
synthcsisinrespirntion 239.247- Sl anaphase 105, 107,3,45,346,347, 348 Badflustburingiensls 485
animal cells 4--5, 15 bacteria 22---3,24,69,431
organisms 24S-SO animals 431.437, 438 antibioticresistant 212- 15, 387
ad ~sion 140-2 domestication of vdld animals 394 bioremediation 450-52
adiposetissue 42 intemaltr.111.sponsystems 151- 70 nitrifying 272
ADP- All'cyde 2.J,6.237 mammals see mammals tran.sfonned 459---{i()
odrenalglands 328 osmosisin 89 balanced polymorphism 389
ad n,nalin 394-S respiratorysystems 172-80 balancingselecrion 388---9
aerench;matissue 255, 2<;6 Slt!etJlsounderindiddu,a/,anim,a/s barleyfruit 339
oerobkrespirntion 172. 242---{i, antagonism 337 baseadditionmutation 367
251,254---S antibioticresistance 212- 15,387 basedeletionmutation 367
Africanelephants 413 antibioticresistanccgenes 460,461 basesubstitutionmutation 367,368
agricu.lrure Z76,443 antibiotics 210-12 BCGvacdne 201- 2
,.,)-.,
adapationsofcropplants 255-7, antibodies 221-4 , 225
monoclonal 226---3()
"""'%380
behaYiournlseparntion 406----8"
genetically modified organisms anticodoru 122,125,126 belttransect 423,424
485--9 antidiuretic hormone (ADH) 3()}-5 Benedidstest 30,32-4
Agroruclerium 487 antigen-antibodyreacrion 220-(225 benigntumours 104
A,grosristetmis(Bern grass) 4()4 antigenreceptors 221 Bentgrnss 404
air composition in the lungs 180 antigens 204,219,232 beta-carotene 486
airspaces 130 aorta 154,155.1S6 betacells 293.295
albinism 369--70 apoplastpathways 137,138 ~ glucose 31
aquaticuniceUularanimals ll9 ~sheets 47
akoholicfermemation 253.2S4.2S6 archaea(archaebacteria) 433 biruspidvalves 164,165
aldoses 32 arrhythmia 170 bilayer,lipid 75,76,n,90
alienspecies 446---9 aneries 154----6 bindingproteins 94
alkalinophiles 431 anerioles l55,1S6 bindingsites 94
allelefrequency 39(),405 anificialimmunity 231 binomialsy.stem 429-30
339-40,342,3'i 3.34S.3~,357 anifkialselection(selectivebreeding) 39}-.8 biodiversity 416-28
510
Index
importance, 442--4 carbondioxide 161.162 chiasmata 345,347
threatsto 439----42 =ncentrationandphotosynthesis chi-,;quarOO(.f)test 364-5
bioinformatics 467- 76,482--4 2,274--S.Z76 Cblamydcmonas 2,434
biologicaldassiftcation 419---39 carbonicanhydrasc 161 chlorophyll 20, 146,261,26}--{,
biologicalcontrolmea.sures 449 carbonmonoxide 161,181,188 chloroplasts 4,5,7,11.15,19-20,
biologicalmolerules 29-53 carboxyhaemoglobin 161,188 25.130
testingfor 19---30 carcinogens 104,105,181 isolated chloroplast suspension
seealsocarbohydrates;Lipids;proteins cardiaccycle 166----8 Z77- 78
bioremedfation 450-52 cardiacmuscle 16}-S photosynthesis 261,266.267,Z76,
bio,ensors 296 cardiacoutput 168 U,-00
bio,phen, 441 cardiovasrular disease (CVD) 1@-70 cholera 194----6,232
biOlechnology 467 carotenoids 26, cholesterol 43,75
bi01icf:i<1ors 3H carriers 358---9,469 cholinergicsynapses 319-21
birthweight 386 canilagerings 175,176 chorionicvillussampling 471,472
biuretteSI 30. 45 . 46 Casparianstrip 131,137,138 chromatids 96.99.106
catabolicreactions S6.S7 chromatin 16,96
"'"""
cirrulationsystem 1Sl---{i3,173
excretion and blood water
catalasc 6L62 chromatography 26}-4
Catalogueofllfe 420 chrom060flles 16,9S-102.342- 3
b:ilancc, 297- 305 catalysts 57---{il meio.,is 101,102,345-8.348-9
bloodcc,Jls 152- 3 seeaiS>Jenzymes mitosis 101,102.105---7,108
n,d 49,152,153.1$4 cattle 394-S '"--Theoryoflnheritance 364
"'hites,;ewhitebloodcc,Us cell---a,llre=gnitionsite 94
bloodclottingrnechanism 371,468 cellcyde 102---3 chronic obstructive pulmonary disease
bloodglucosc,regulation 292- S celldivi.sion 97- 105 (COPD) 182,184
bloodpressure 187,300 uncontrollOO 104--S cigarettesmoking 181--9
B-lymphocytes 221--4. 225 seeaiS>Jme· ·s· 'to.,'s cilia 21,174.182,183
bodytemperature 250 cellelongation 338 ciliatedepithelium 174,180,IS}
homeostasis 289-91 cell-mOOiatOOimmunity 222 cirrulatorysystem 151---{i3
Bohreffe<1 161 cells 1- 27 CITES (Convention on International Trade in
botanicgardens 446 animal 4--S,IS Endangen,dSpecies) 450
brain 312,314 basicunitsoflivingorgani'lms classes 430
broad-spe<1rumantibiotics 210,213 nikaryotic 22.2}-4,Zl,431 classification 429-39
bronchi 173.174-6 plant 4--5,15,132--4 cLimatcchange 410,443
bronchioles 173,174--7 prokaryotic 22- 5,27,431 Clini'lti,i'"' 70,71,296
Brown,Roben 3 rolesofproteinsin SI clottingfactors 468
Blcrops 485-{i size 2.21,Zl,79--8() codingstrand 125, 126
bulbousheads 249,325,326 specialisation 1 codominamaUeles 355,356
bulktransport 78,93 cellsignaUing 77,295 codons 119,120.121,122
bundlesheathcc,Us 281,282.283 cellsurfacc,membrane 4,S, 14, IS, 16, coenzymeA (CoA) 239
by-passvessels 158 ;+-9<
movementacross 7S-94 cohesion-tensiontheory 141
structure 74--7 collagen 48
c,pathway 200,281,282 celltheory 3.4 coll=:insducrs 291'1,302- 3,30}-S
C,plants 28}-1 cellularrespiration 2}1----59 collenchymacc,Jls 133,134
calorimeter 240 cellulose 37 colourbLindness 359
C.Mncycle 270-1 cellultrastructure 12---14.14--25 combinedoralcontracep1:ivepill 332
cancer 229 cellwaU 4,S, 15, 20,211 combustion 235
SIi
Index
compo,rndlightmicrosrope 2, 6--11 andthegene1kcode 118-19 clectronmicrosc~ 4.12- 14
concep1ion 332 mitochondrial(mtDNA) 402---3 electrontransportchain 239, 243.246,247---9
condensationreacrion.s 35,SO,lll inproteinsynthe-sis 120 electrophore-sis 464---5
conne<1ingcorridors 444 replication 98, llS-17 embryo 332,471- 2
-=tion439--S2 sateUiteDNA 483 embryobiopsy 481
new developments 447- Sl descendinglimbOoopofHenle) 302---3 enibryoselection 481
-=tiven,plicatiooll6 diabetes 295 entphysema 182
<nntinuous varfation 3S3. 3n,378-9 diaphragm 173,177 emulsiontest 30,41
=ntracep1:ion 332 diastole 166---{;7 endangeredspedes 441
=ntracep1:ivepill 332 diastolkpressure 187 prolection 444----6
contractilevaruole 2,10,S9 differentialrespirometer 258-9 endemicdisease 193
=ntrolandco--0rdination 311-40 diffusion 78,79--90,128 endocrineglands 328,329
mammals 311- 33 digitalmicroscopy 8 endocrinesystem 291,303. 328-33
plants 311,334-40 dihybridcross 360-63 endocytosis 93
Convention on lmemational Trade in diploid cells/organisms 101,102, endodennis 131,137,136
EndangeredSpecies(CITE.5) 4SO }42-3. 357 endometrium 329--31
convolutedrubules 296,299.301.303 dipoles 36-7,51 endoplasmicn,tirulum 13,lS,16--17
coordiruitor 286 dip.sticks 296 =•qzy
coraln,efs 410,442 directionalselection 3ff7 controlledtransferinrespiration 234-6
cormorant 417- 18 Din,<10bservation,Treaunent.Shortcourse ene'8Yvaluesofn,spiratory
coronaryaJteries 163 (DOTS) approach 203 substrates 240
corpuslmcum 330,331 disaccilllrides 35---{; fromfatsandoils 42
crntex(kidney) 298,29') disrontinuousvariation 353,377,378 photosynthesisasenergytransfer 261- 72
counter-eurrentmultiplier 302 disease 192 andn,spiration 234-S9
<UValentbonding 36,Sl enhancersite 374
cows 394-S r:r:~:~:-::~~a:~~s':ase
Crick,Fraocis 114 non-infectious 192 binttmediation 4S0-2
criticalvaluesforthet-test 382 disrup1iveselection 388----9 effectonphenotype 366,360
cropplantssoeagrirulture distal=nvolutedruhule 299,303 environmentalfa<10l"Sandpopulation
Cf06Singover }iS,346,347,348. 349,400 disruJbingfactor.i 391- 2 size 386
cryophiles 431 disulfidebonds 47 Hmitingfacrorontherateof
crystaljellytish 463 DNAlig:,se 115, 116.455,4'>8 photosynthe-sis 273---{i
cyclic AMP (cAMP) 294 DNAmicroarrays 465-6 enzymes 48, 56--71,94
cydicphO!ophosphmylation 271- 72 DNApolymerase 11S-16,45S,4s6--7 concentration 67
cysticfibrosi.s 469-70,471,HS . 476 DNAprobes 464,471,482 facrorsaffecringacrionof 6}-7
cytokinc,sis 101,102,106,107.}iS DNAproflling 482--4 inhibitorsof 69---9
cytoplasm 2,4,5,7 DNAsequencing 455,464 modedaction 56---{,3
cytosine 110,111,112,113.118 DNAYiruses 25,439 operationof 62--{;
cytosol 14 dodos 414 rateofreaction S7--{;1
Doll, Richard 186 specifidtyof 59
domains 431-3 epidemics 193
dairyca!de 394-S domesticationofvdldanimals 394 epi<kmiology 184
dandelion 380 dominantalleles 352,357,3@.370.469 epidermis 7,130.1 33,139
Darwin,Charles 393. 39') doubledrrulation 152.155 epithelium 174
double-erosshybridisation 398 ciliated 174,183
deCOntpo6efS 417 doublehelix 112,113,114 Escbericbiacoli(E.col,) 2.22- 3
degreesdfreedom 365,381- 2 drought 143.283,30') gen,etkengineeringofforinsulin
DEI.L\pro1eins 375 dwarfrnrietie-s 397 production 4S5-{;2
demes 409 estcrbonds 40
<knaruringofproteins S0.64.65 ethics
dendrites 312,313 ecologicalseparation 4()6....g genetechnology 478-9,481,489
dendroru 312,313 e=systems 417,420 gcnctkscreening 479
<koxyribonuclekacid(DNA) 98. 100,110, ecotouri.sm 443 eukaryotes (eukaryota) 433.438
112-13,114,llS-16 edgeeffect 444 generegulation 374
complementa,y (cDNA) 4S6. 465-6 effector 288.311.314 gcncticmodific~tionof 462
damageduetosmoking 185 eggcells 329--31 eukaryoticcells 22,2}-4.27,431
DNAprobes 464 .471.482 elasticconnectivetissue 178,179 evaporation S2- 3
DNAprofiling 482--4 electricalinsulation 42 evolution 399-413
DNAsequendng 455,464 electrocardiography 1@---70 molecular evidence for evolutionary
genemutations 124----6 electroncarrierproteins 94 n,lationships 400-3
Sil
Index
exdtatorysynapses 319--21 genetechnology 4S4----89 graincroJlll 395---,8
excitedelectrons 168---9 applied!omedidne 467---fl4 grana 20. 267,260),279,280
excretion 297---306 principlesof 454-{i6 grasses 388
exocytosis 93 social and ethical aspects gratirule 9
expiration 177,178 477---fll,4@ greenfluorescentprotein(GFP)gene 463
expressionvector 460 genctherapy 46S----9. 47s-.81 greysquirrel 448
extinction 409--H genetically-modifledoqpnisms growth movements 334-5
extrncellularenzymes 57 (GMOs) 485-9 growthregulators 335---40
extremophiles 432,432,438 geneticcode 110,118-26 guanine 111,112,113.118.119
eyepieceleru 6 gencticcounselling 473 guardcells 139,307---fl
genetic crosses
dihybrid 360-3
fadlimeddiffusion 81- 2, 316 monohrbrid 3SQ----9 habitats 417,418,420
factorVHI 468 probabilityin 364-5
families 430 geneticdictioruiry 118.122,124 restorationofdegrnded 450-2
familypedigrec, 36S-70,473,474 gencticdisorders 359,479 stability 440
fats 39--"13,239 cys!icfibrosis 4@-70,471,475.476 hacmgroup 49
see also lipids haemophilia 371,468,474 haemoglobin 49, 160--2. 173,401
fattyacids 39,40,239 screeningfor 4@---74,479 haemoglobinicadd 161
ferns 436 sicklecellanaemiaseesicklecell haemophilia 371. 468,474
fertilisa!ion }4}-4,3SO anaemia/trait halophytes 431
in-dlro 447,479---61 geneticdiversity 442- 3 handlens 6
fibres 132,133 gencticdrif! 392 hap[oidceUs/organisrns 101,102,343,357
fibrin 48 geneticenginec,ring 454--5 Hardy- Weinbergformula 390---3
fibrouspro!eins 48 health
finches 407---fl :r:i=:bi~;~~;:.uman insulin smokingand 181---9
flaccidity 84,85,86. 88 production 455---{;2 se<ia/sodisease;infectiousdisease
flaseHa 21 markcrsfor 462---3 bean 15(16}-70
flavineadeninedinucleotide(FAD) geneticfingerpriming 455.464.482--1 heanblock 169.170
238,239 genetics 378-2 he~n ra!e 168
floweringplams 436 inheritance },2-74 heavymetalion!olcrance 404
cornmunicationandcontrol 334---40 geneticscreening 4$---74,479
fluid lllOllaic model of membrane geneticvariation 348-9,377---fl3.400.40L hcmkclluloses 39
structure 76.77
fluorescemmarl<ers 463
folliclestimula!inghormone(FSH) 329---31
"'
genomes 372,405,467
genomics 467
hepaticportalvcin 154
herbidde-resistantcrop;s
herbicide,, 451
487---9
513
Index
human population growth 333, ~40---1 lmemational Union for the Conservation lignin 132
humoralimmunity 222 ofNature([UCN)RedLis!oflllreatened limitingfactors 27}-78
Huntington'sdisease 370 Species 4H llncolnlndex 425, 426
hybridomacclls 228 imerphase 102---3 linetranse<1 423
hybridvigour 396 interspedficcompetition 419 linkreaction 243-S
hydrogenbonds 36, 47,51,52,82 imerventionstudies lSS lipidbilaye r 75,76,82. 90
hydrolysisreaction 35 , SO intracellular enzymes S7 lipids 30,39----43
hydrophilicproperties 37,42,53,75 intraspeciflccompetition 419 respiratorysubstrates 239. 240
hydrophobkpropenies 37,42. 53,75 introns 463 lipo60mes 47S,476
hyperglycaemi.i 292 invasivespecies 440,----9 liverceUs 294---S
hypenonicsolutions 86 in-drroknilisa!ion([VF) 447,480--81 'lockandkey'hypothesis S9
hypillle 435 iodine!est 3(),38 torus 3"2.343,3'xi,3S7
hypoglycaemia 292 ionchanncls 317- 18 loop of Henle 298, 29'), 302
hypothalamus 290-1,303,304, 328, ionconcentrationregulation 303 lungcancer 1S2,183-S
329, 330 ionicbonds 47 lungs
hypotonicsolutions 86 ioniccompounds S3 gaseousexchangc 173-SO
transponolrespiratorygases 160----3
irre,,ersibleinhibitors 69 luteinising hormone (LH) 329-31
i<kmiftcationusingDNAprdi.Ling isletsollangerhans 293.294 lymphatic system 1S9, 160
4S3,48'l isotonicsolutions 86 lymphcapillaries 158--{iO
iguanaLizards 406--7 ivorypoaching 413 lymphducts(lymphatics) 1S9, 160
immobilised enzymes 70-1 lymphnodes 160
immuneresponse 219--24,225 J lymphocytes 153,219-25
immuncsystem 218-25 Japaneseknotwred 449 antigen-antibodyreacrion 221-l . 225
immunity 218-32 B-lymphocytes 221-l . 22S
amibodies andvaccination 226---32 T -Jymphocytes 204----06, 221-l, 22S
immunocompromised patients 205 lysosomes lS,19
immunologicalstudies 401- 2 ketones 305 lysozyme 48
impulses 312. 31S...-16 ketoses 32
inbreedingdepression 389 kidneys 297- 306
independemassonmem }iS---9,360, kinases 295 macromolecules 36,SO, 110
361,400 kingdoms 430,431,434---37 soealsopolysaccharides;proteins
indoleaceticadd([AA) 336- 337 knotweed 449 macrophages 19,93, 178,179,219,220
inducedfithypo(hesis 60--l Krebscycle 24}-5 222,223
inducibleenzymes 373 magnesiumions 144,146
indusufalcatalysts 69--71 magnification 10-11
indusufalsolvents 452 laboratory-based investigations magpies 429
infammortality 386 Lack. Da,id 407-8 maize 261.282
infecriousdisease 192- 215 /acoperon 37}-4,460 Btmaize 48H
amibioticsandtreatmeruof 210-15 Jactase 61 selectivebreeding 393----8
casesrudies 194-210 la<1kacidfennentation 253.254 major histocompatibiLi!y complex
inflammation and immunity 218-32 lacrose 36 (MHC) 220-1
infertility 479---fll la!entheatofvaporisation S2---3 majormineralelements 145
inflammation 21S.-19, 220 UlwoflndependemAssonmem 360 malaria 197---9,232,389. 467
inhibi!ors 66---9 UlwofSegregation 352 malecirrumcision '207
inhibitorysynapses 321 lea,·es 130 malignam rumours 104
inorgankions 144----6 waterstress 309 Malpigltianbody 298.m
insecticides 4S1 leeuwenhoek,Anthonyvan 3 maltose 36
insect-resistantcrops 48H leucoq1esse<1whi!ebloodceUs
inspiration 177,178 leu=vlasts 20,38 controlandco-ordination 311- 33
insulin 293,294,295,468 Jifeexpectancy 333 homeostasis 206---306
productionbygeneticengineeringol ligase 115,116.4<;8 internal transpon 151- 70
E<cbericbi<acoli45H2 light 262 markers 462---3
integralproteins 76 intensityasalimitingfactorin mark,releaseandrecapture(MRR)
interoilmeddiscs 16}-4 photosynthesis 274---5.276 technique 425-26
inten:oswlmuscles 173,177 light-dependent stage of massextincrionevents 409
intemaltransponsystems photosynthesis 266-70 mass flow 129, 147
heanasapump 16}-70 light-independent stage of Mauritius 41}-14
mammals 151- 70 photosynthesis 266, 270-2 measles 208---9,232
plants l;is....l9 lightmicroscopy 2,6--11,108 m<."Chanorereptors 315
514
Index
medicine 467- S&l M),cobacrerium1ubercu/asis 200 see,a/solipids
medulla 296,m.302 myelinsheath 313,319 oilspills 4S0-1
meiosis 101- 2,3,42---49 mrOObrils 322,323 oogenesis }t4
melanin 379 myosinfllaments 249-S0,324 operatorgene 373,374
membranepro1eiru 76,94 mrxornatosis 385,449 oralrehydrationpack 195,196
membranetransponmechanism.s 78-93 orders 430
memorycclls 223,224 organelles 4,25
Mendel.Gregor 339. 3S0,3S1,364 NAD(nicotinamideadenine suucrureandfunction 14--22
menstrualcycle 329-31 dinudeotide) 237,236. 267 organicacids 308
Merlin 427 NADP(nirotinamideadeninedinudeotide organic compounds 29-53.S7.S8. 262
mesophyllccUs 130,281,281 phosphate) 266,267 orgarusms
mesophytes 142 ccllsasthebasicunitsci 14--TJ
messenger RNA ( mRNA) 113, 120, 124 naruralimmuniry 224,231 classiflcatlonof 429-38
4S6---7,463 naruralselecrion 38}-93 osmoregulation 296,303-5
metabolic,..is!e 297- 305 evolutionby 399---400 O'lffiO'liS 78,82---9()
metabolic,..i!er 42 narurereserves 444--S ovariancyde 329-31
metabolism 14,S6--7 negativecorrelatlon 426 ovaries 328,329-31
metabolites 56 negatlvefeedback 288,306,330 overpopulation 333
metaphase 105.107.3,4S.346,347,348 Neo-Darwinism 400 oxidativephosphorylatlon 243,246
metastasis 104,183 nephrons 296.299 oxygentranspon 160---1.162- 3
nervoussystem 288,290,291,312- 22 oxygendissociationrurve 160,161
Michaeli'l-Mentenconstant 67 neuromuscu.larjunctions 324---S oxyhaemoglobin 160,161,162
microhabi!at 417,418 neurones 312- 22
microme!er(micron) 2,4 neutrophils 219,220
microscopy 1- 14,108 newborn scn,ening 472 Padniancoipusde 315
ele<1ronmic!06COpy i, 12- 14 niches 417- 18 paddyf\elds 255-7
lightmicrosropy 2,6-11.108 p;,ncreas 293.294,328"
microrubules 19 (NAD) 237, 238. 267 pandemicdisease 193
migration 393 nicotinamideadeninedinucleotide Paramecium 411,434
Mimosa pudic,a 33S phllllphate(NADP) 266,267 p;,rasites 467
mineraldeflcieocyinplams 145---{i nicotine 181,1S6 parenchymacells 133,134
mineral elements, major 14S seea/5vsrnoking partialpermeability 82,83
mitod!ondria 1S,18,2S, 247,248.2Sl.322 nitrmeions 144,14S partial pressure ~9, 161. 162, 100
mitod!ondrial DNA (mtDNA) 402- 3 ni!rifyingbacreria 272 passiveimmunity 208,231
mitosis 99,101----8,343,3,44 norlesciRanv:ier 313.314,319 passivesmoking 182
chromosome behaviour 105---,8 non-eompe1itiveinhibitors 68,{:f) Pas!eur.louis 3
~™Rvaccine 208,2(1) non-qclicphotophosphorylatlon U:l).T/2 paternity 484
molecules 26,27 non-governmo,ntaloq;;anisations 4SO pathogens 193
biological 29-S3 non-infecriousdiseases 192 peaplants 339-40. 3so-.J,36o-2
monodonalantibodies 226---30 non-regula!ors 287 Pearson'slinearcorrelatlon 426,427
monohrbridcr05ll 3SO-S9 non-relfpro1:eins 204,219,221,224 pectin, 39
monolayer,lipid 75 non-venebrmeanimals 437 pentoses 34
monomers 36. SO noradn,nalin 319 peptidebonds 4S,46,S0
monosaccharides 31- (36,SO nucleardivision 97- 105 pericardium 163
Morgan, Thomas 364 meiosis 101- 2,34}-9 pericycle 131
morningafterpill 333 mitosis 99.101----8.3'l3,3,4S periphernlnervowsystem 312
mosquitoes 197--99 nuclearenvelope lS,16 periphernlproteiru 76
mosses 436 nucleic adds SO, 110-17 pennaneruvacuole lS,20
motorneurones 312,313 nucleolus 15,16 pesticides 451
MR.'iA (med!idllin resistant SiapbJ•lococcus nucleotides 22,110-12,238-9 petrification 440
m,reus) 212.213,387 nucleotldesequencc 368
multicellular organisms 2, 97
multldrug-resi'liam TB (MOR-TB) 201, 202
DNAsequencing 455.464
nucleus 4.5,7----8,15,16. 97----8
'" cffectonenzymes 65
blood 303
multiplealleles 356 nullhypothesis 381 pha8'X}1e~ 153. 219
mwdecontraction 249-S0,322- 27 pha8'X}10Sis 93,219
mutagens 367 phenotype 350-71
mutations 124--S,367- 71,393 objecrivelens 6 cffectofenvironmo,m 366,360
andcancer 104--S oedema 160 cffectofmutatlons 367--41
genetlcdisorders 469,471 oestrogen 329----31 phenrlketonuri.i(PKU) 472
myastheniagrav:is 224 oils 39-H phloem 129.132,133,146----9
SIS
lnde,
phloemtt:lnspo!t
,_,,,
p!Kl6pholipids
1"47-9
39,H-3, 74--S
poooopOOSphorylotion Uil,Uf),271-2
pholoreccptors 315
phomresplra1lon ?8),281
photo.1yn1hcsls 130.261---fl3
adaplatiorufor 279413
-
prey--ptt<latorpopulationOfidllation
positivccorn,latlon 426
posHygotlc rcprodoclivcisolation
pconiumchannels 317-18
pcossiumions 317
»dium-poo,ssium ion pump 92
po(omctcr 141,142
1(19
Q
quadr:1.1:1 421,422
quatcnuryinsulinprodoction
rabbic,; 384----S,449
460-1
quatcmary:1trucrurcofapro1cin 49,226
rainforcstdestruction 412,441-2
asencrgytr:m~er 261- 72 prcgnancytcsting 229,230 random assonmem }16-9, 360. 361. 400
environmcntondr:m,of 27}-6 prcnatal...::rttning 471- 2 r:andomfu,ion 348,400
timitinsfactor5 Zl}-78 pl'C'SSIJl'Cflowhypoihesis 148.119 r:andom sampling 421, 422
process 266-72 prcssurcpotential 8'1----S rc:alw>rption 301.303--06
phou)systcms 261-69 preventive health l"C'«J)IOIS n,9'i,298.306,j l4
phylum 130 CVD ,.,_. s,m,sory 31S
-
pigeoov:uictics 393 lnf«1iousdiscase 193.196.198-9. rcccssivc,allcic,; JS2.J57.3$-70.469
pillOC')'006is 93 201-2.206--7,208 rccomhinantDNA 4',1
piruii:uyglalld 303-5,320,J28,330 pl'C')'--f)t"Cdtorpopulationoo:cillalion ~nantplasmidli159-6)
pl~r11.3 332 rccomhinanC!l 348,360
plandiagrams(tissue maps) 134 p rc-zygoticrcproducth'eisolation 1(19 rC'dbloodcells 49,SO,IS2.
plank1on 384 primarypumps 147, 146 153,154
plamcells 4-5.IS.130, 133,134.136-40 prima,ystnicturcofpro1cins 45. red,r=ncolourblindncss 359
plamgrowlhrcgulators 33S,-4() 48.226 RcdListof11Ircatcw,dSpcdcs 44 1
plants 43L.·H6.438 prol»bility 364--S rcdm:rcactions 237,U:,J.
-~""
rontroland<:<>-<>rdination 3}4----40 produccn 417 rcducingrug,tr5 30,32-4
cropplanl5seeagriculrure protllctranllttl 423.424 rcfle:xarcs 314
flowcri1133~0,-'\36 proscstcronc j29---31 rcfr:actory~l(,6,318
homeoMllsis 307-9 prok:uyofa 37}-4,431,4JJ. 438
imemaltranspon 123-49 prol<:uyoliccc:lls 22- S,Zl,'131 rcgul:Uorygcncs 373
06ltl0a5in~ prophasc 105, 107,345--6.}'18 rcby n,,urooes 312,313,314
phol06)'fllhesisSNphorosyn~sl!i pro,ct>ctlcgroup 49 rcnalcapsulc 298,299-300
plasnu IS2.1S3. 1s-4 pro1cinflwrt=c,; 324,325-7 rcplkalionofDNA 198, 115-17,211
plasnucell5 219.223 proicinpump, 90--2.94. 144 rcplkatlonforks 115
plasmids prolcins 3(),44--51 . 305 rcprcssibleemymC11 373
R-plasmids 460.461 lnblood IS2 reprcsaormolcrule 37}-4
Tiplasmid 467 channclproicins 81---3,94 reproductivcisolatlon 406.409
vectorforinw~ngcnc 4S&-(,O tknaruringof S0,6S rcsolutlon(rcsolvingpowcr) ll
plasmodesm:ua 20,U7 functionsolmembran,,protciru respiration 2}'1-59
Plasmodium 197-8, 19),467 76.94 :acrobicl72,24M2.~.262
pbsmoly&is 85.86.ffl-8 humanpro(rinsandtrc:aunc,111of :uucrobic 2S}-S7
plastids 20.38 disease '468-9 rnffrollcdtnn,;fcrofcncrgy 2}'1-7
platdeu IS).154 rC9piratorysubstrat"5 23'),240 =pirarorygn=pon 160-1
rolcsof SI =piraroryquoticnt(RQ) 241
'""""'"
poi5005 $ JOO respiratorysurbcc,i 173
polarmolec\Llc:I 36,Sl,S2,90 pro!cinleqUNtOedata400--I rucpirometcr 2S7-9
polygcnicinheritance 378-9 prolcinsynthc.'lis llS-26.211 m;tlngpotentlal 316
polymerascchainrcaction(PCR) stages 120-2 m;trictioncnzyme(rc•triction
·457,483 prolcome 4'>1 cndonoclca..,) 4S5,458.459.464
polymers 50. 110 prolcornk• 467
polynudCO(idcs 112 proloctisU 431,4.}t,438 revc....., transcripUse
455.456
see1llsonuclcicacids prolonpumps 337,338 R-gnJUp., 32,44----S
polypeptides 4S.l1S prolO·O<U;Qf!Cr>CS IQ,I ribonu~icadd(RNA) 110.113
seetllsoprotcins proxinu!convolmedrubuk m.301 me"1lCfl8C"RNA(mRNA) 113.120. 123,
polysaccharides 36-9,50,76 pulmonaryancry 178.179 124,4S6---7,-463
polysomc 122 pulmon;uyrirrubtion IS4 ribo&omal 432
polyuns:uuratcdf3ts 41 pulmona,yvcin 178,179 tran.sfcrRNA(tRNA) 113, 121 .. 123
populaliongcnefics 389----393 pulse 1(,6 riboMlmaJRNA 432
populalioru 389,419 pump proteins 90--2.94, I« ~ l S . 16,17, 18, 120,121,122
HUC11.1ationofpb11ktonpopul:Kion• 384 Punn,,ngid 3S3 ribul,...,biphosJ»la(c(RuBP) 270,271
humanpopulationg"""'lh pyraOORring 31 ""'YJ6
m.,wo..1
"'
Index
swampplants 255- 7 smoothendoplasmicn,tirulum(SER) synap1icdefi 319,320
vitaminAenhanced 486---7 15.16.17 synergism 337
RNA polymerase 120 smoothmuscle 175,176 systemiccinculation 154
RNAviruses 25,429 sodiumchannels 317- 18 systole 166-7
roothairs 131,13S-7 sodium-potassiumionpump 92 systolicpres.sure 187
roots 131 soils.bioremedi.itionof 451- 2
roughendoplasmkn,tirulum(RER) solutcpotential &l.85
15,16,17 solvents tachycardia 169,170
R-plasrnids 460,461 industrial 452 targetcells 329
rubisco 270,280.261 water as a solvem 53 targetorgans 328
somaticgcnetherapy 475,476 taxonomy 429---39
sorghum 143.281,283 telomeres 101
sourceareas 147, 148 telophase 111 , 105,107,345,346,348
saltatO<yconduction 319 Southemblooing 482 tempera run,
sarcolemma 163,164.322 Spearman'srankcorrelation 426,427 bodytemperarun, 2S0,289-91
sarcomen.-s 324 specialisationofcells 6 andrnteofphotosynthesis 275---{,
sarcoplasmkretirulum 322.323 speciation 386-9,403-9 andrnceofn,actionofenzymes 6}--5
sateUiteDNA 483 species 403-4,416--17,429-30 andrnteofn,spiration 252,257--9
saruratedtriglycerides 41 - 2 alienandin,11sive 448-9 temperarun,coefficient 65
scalebars 9,10 biodiversity see biodiversity tertiarystructureofaprotein 47,48,226
scanningeleCTronmicroscopy 13 genetkdiversityandevolutionofnew testcross 357
Schleiden.Manhias 3 species 442- 3 dihybrid 362
Schwann,Theodor 3 Spedes2000programme 420 monohybrid 354
Schwanncells 313 specifichemcapacity 52 testes 326
secondary pumps 147,148 specificity o f enzymes 59-4> testisdeterminingfactor(Il)F) 357-6
secondarystructun,ofaprOlein sperm }t3-4 testosterone 329
47, 48,226 spermatogene-sis 344 testsforbiologicalmolccule-s 29---30
secondmessenger 294 sphygmomanometer 187 lipids 41
seedbanks 44{, spin.alcord 312 non-reducingsugars 35
selection 383----96 spindle 19,105,106, 107, 346,347,3,48 proleins 45. 46
aniftcial(selectivebn,eding) 393-S spon,s 435 reducingsugars 32-4
natural 383-93.39')-400 stabilisingselection 386 starch 38
selectivepn,dation 392 stagemicrometer 9 therapeutic ~bonions 481
semi-conservativen,plication 116--17 staining 464.4155 thermopltile-s 431
semilunarvalves 164,165 starch 6. 30. 38 thermon,ceptors 315
sensecells 315 statisticaltests 380-2. 426 thermon,gulation 189-91
sensoryneurones 312,313 Stele 131 thickfilaments 324
serum 401 stem 129 thinfilaments 324
seven,combinedimmunodeflciency stemelongation 339-40 thinse<1ions 12.13
(SCJD) 4@ steroids 43 thorax 173,177
sex chromosomes 357--9 stimuli 312,318 thresholdofstimulation 318
sexdetermin.ation 357~ stomata 130, 139-40, 307--9 thylakoidmembranes 20,267,268. 269,
sex-Linkedcharn<1eristk:s 3':>8 openingandclosingof 308---9 V,,,00
sexualn,producrion 343-4,3SO striatedmusde 322- 7 thymine 110,lll,112,113,118,119
shag 417- 18 strokevolume 168 thyroidgland 326
skklecellanaemi.i/trait SO, 125,126.232 stroma 269,270. 'Z79. 280 thyroxin 291
355,368,388-9 strucruralgene-s 373 Tiplasmid 487
n,sistancetomalaria 196.389 subcutaneousfat 42 tissuefluid 157--4:l
sieveplate 146,147 sub-species(varietie-s) 404 . .06 tissuemaps(plandi.igrams) 134
sieverubes 133,146,147,141!149 substrate 59 tissues,exchangein 156-60
Simpson's Diversitylndex 427 concentration 66 T-lymphocytes (CDt helper cells) 204----6,
sinkan,as 147,148,149 sucro,;e 35---{i 221-4 . 225
sinoatrialnode(SAnode)(pacemaker) 168 sugars 30, 31--9 tonoplast 20
sizen,lationships 26---7 surfacean,a : volume ratio 79---,61 trachea 17~
skeletal muscle 322- 7 surfactant cells 178,179 transcription 120,211
skin 289-90 'survivalofthetlnest' 400 transcriptionfa<1ors 374--5
sliding-filamenthypothesis 32S-7 sympatricsp,,ciation 408 transects 42}-4
smallpox 210,232 symplast pathways 137, 138 transfercells 147,148
smoking 181--9 syn.apses 319-22 transferRNA(tRNA) 113,121
517
Index
transfonnedbacreria 459--4) =ntractile 89 propeniesof 52---3
transgenicoq;anisms 4S4 permanent 15.20 waterpcuential 83-4,85, 86--7, 136
transl3tion 121- 2,123 waterpotentialgradient 83
transJocation 146----9 bean 164----6 water-prOOO.ng 42
transmission electron veins 156 Wa!SOll,James 114
micrograph.s(fH,I) 13 vanderWaalsforces 57 waxes 43
transrnittersubstances 319--21 vari3blcnumbertandemsrepeats wheat 396-7
transpiration 139--42,307 (VNfRs) 483 whitebloodcellsOeucocytes) 153,154,
transpirationstream 14()-41 vari3tion 348-9, 377-83,400, 401. 420 219,224
varieties(sub-species) 404. 417 lymphocytessrelymphocytes
internalseeimemaltr:msponsystems vasarecta 302- 3 phagocytes 153.219
membrane transpon mf'Chanisms vascularbundles 129,130 Wilkins,Maurice 114
vasculartissue 129,130 wilting 88, 307
tricuspidvalve-s 164,165 World Health Oqpnisation (WHO) 203
triglycerides 39--42.240 forgenetherapy 475 World Wide Fund for Natun, (WWF) 450
triosephosphate 270-1 transterringdisease 197
trioses 34 veins 154-{i
tripletrodes 118,119.122---4 venacava 154.155 Xchromosome 357- 59
tropicalpl3nts 280-3 ventilation mechanism 173,177 xeromorphicfeatures 143
tropomyosin 325, 326 ventricles 164.165 xerophytes 142--3
troponin 325,326 ventriculardiastole 166,166 xylem 129,132
r-test 360-2 ventricularfibrillation 169.170 xylemvessels 132
ruberrulosis(fB) 100-3.232 ventricularsystole 166,167, 168
tumour-<1ssociatedantigeru(fAA) 229 venules 155,156
tumours 104. 183 Venusflytrap 337
v,, _ _ 357---8
tumom.suppressinggenes 104 venebrmeanimals 437 yeast 252,435,462
tuq;idity 84.85,86.88 vesides 93 yield 397,485-7
ruq;orpressure 307,308 V.briocbo/eme 194
Victoria,Queen 474
Virchow,Rudolf 3 Zlines 324
ultral\Jtrntion 29.}---300 viruses 25---{\l.7, 212.438-9.475 zoological gardcns(zoos) 445
unkellularoqpnisms 1.2,97 vitaminA e nhanredrice 486--7 zygoie 350
aquatic 89
unsaturatedtriglrcerides 41- 2
uracil 110,111,113 Wakcfield,Andrew 209
urea '1!)7,296 Wallace,Alfn,dRussel 399
urine 29S-305 water 36,51- 3, 90
analysi.'lindiagnosis 305 -=tioninlheloopofHenle
uterioecyde 329-31 30'-ol
excretion and blood water
balance 297---306
vaccination 201 - 2,206,208,209 lossduringgasexchange 175,176
210,231- 2 metabolic 42
vacuolc,s 4,5 movementthroughaplam 13s--43
518
Acknowledgements
The publishers would like co thank the foUowing for permission to library: p.183 e Mo"'11un Animal Health Ltd/Science Photo Library;
reproduce copyright material p.185 Cl AP/Pn,ss Association Images; p.lS8 Cl Ulet lfansasti/Geny
Im ages; p.194 Cl Dr. Hans Ackermann/Visuals Unlimited. Inc./
Phom credics : Science Photo Library; p.196 I e Science & Sodety/SuperStock, r
p.5 1 ClJ.C. REVY. ISM/SCIENCE PHOTO LIBRARY. r Cl Gene Cox: Cl Paula Bronstein/Geny Images; p.200 r Cl James CavaUini/Photo
p.S e Gene Cox; p.10 Cl Power and Syred/Sdence PhOlo library: Researchers. lnc./Science Phom library, me Adam Han-Davis/
p.11 /ClBiophotoAssocfates/SdencePho!Ollbrary. r Cl J.C. Revy, SciencePhotoLibrary,b Cl BSIPSA/Alamy; p.201 /ClBiopho!oAs--
lSM/Science Photo library; p.12 Cl Sinclair Stammers/Science Photo soda!es,IScieoce Phom Llbra,y, m Cl Dr P. 1,farazzi/Sdencc, Photo
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Aberdeen University. m Cl Eye O f Science/Science Photo Library. Ncmir/Sdence Phoco Libruy: p.210 e Medical-on-Line/Alamy;
b Cl Power and Syred/Sdence Photo Library; p.14 1 Cl Phototake p.214 I Cl RGB Vemures UC dba SuperS!ock/Alamy. r Cl Paul Gun-
lnc./Alamy. r Cl Biophoto AssodatN/Sdence Photo Library; p.1 7 te ning!Science Photo Library; p.220 e Gene Cox; p .227 Cl Science
Medimagc!Sdo,na, Photo Library. b Cl Omikron/Photo Researchers, Photo Library/Alamy: p .235 I Cl ~fanyn F. Oi.illmaid/Scio,nce Photo
lnc./Science Photo Library; p.18 t e Carolina Biological Supply. Co/ Library, r Cl LIU JIN/AFP/Geny lmage-s; p.249 Cl Medical-on- Line/
v-.suals Unlimi!ed, lnc./Sdena, Photo Library. b e CNRVSdence Alamy: p.251 Cl Biophoto Assoda!es/Science Photo Library; p.256
Photo Library; p.19 Cl Don W. Fawcett/Science Photo Library: p.20 r Cl jjayo - FotoLi a, m Cl Dr Keith Wheeler/Science Photo Library, b
t e Dr Kari l.ounauruia/Sdena, Photo Library/ Getty lmage-s. b Cl Dr Cl CJ Clegg; p.279 Cl Dr Kenneth r. Miller/Science Photo Library
Jeremy Burgess/Science Photo Library; p.23 Cl Kwangshin Kim/ p.282 Cl Dr. Ken Wagno,r/Visuals Unlimited, lnc./Sdence Photo
Science Photo Library; p.26 rl e A. Dowseu, Health Protection Library: p.283 e June Green/ Alamy; p.287 rl Cl Silver - FotoLia, ml Cl
J\sency/Science Photo Library, bi Cl Nigel CanLin/Alamy, br e Nigel Kevin Schafer/Corbi.s, mr Cl Jamhonjackson - FotoLia, bi Cl Jan-Dirk
CauLin/Alamy; p.33 both e Andrew Lunben Photography/Science, Hansen - Fotolia, br Cl Juniors Bildarchiv GmbH/Alamy: p.2S8 Cl
Photo Library; p.37 Cl Biophoto Associa!es/Science Photo Library Nickel Electroltd; p.293 Cl Astrid & Hanns-Frieder Michler/Sdencc
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p.39 tCl Phototake lnc./Alamy, bi Cl Bon Appeti!/Alamy, bm Cl loy/Sdence Photo Library. b Cl Garo/Ph.anie/Rex Feature-a; p.299
quaysido, - Fotolia. br Cl Profotokris - Fotolia: p.4I CTrl"vorClifford Cl agefotostock/SuperS!Ock; p.300 Cl SteveGochmeissner/Science
Photography/Sdo,na, Photo Library: p.42 e Science Photo Library/ Photo Library/ Getty lmage-s: p.320 Cl Prd S. Cinti/Science Photo
Alamy: p.48 Cl Sieve Gochmei.ssner/Science Photo Library: p.71 Cl Library: p.323 I Cl Geno, Cox, b Cl Mediocan. University d Aberdeen:
Cordelia Molloy/Science Photo Library; p .77 both e Don W. Fawcett/ p.324 Cl Marl< Rothery, A level Biology, www.mrothery.co.uk; p. 326
Science Photo Library; p.S8 both Cl Biophoto Associa!e-s/Scio,nce Cl Mark Rothery, A Lo,vel Biology. www.mrotllery.co.uk; p.332 Cl cris-
Photo Library; p.100 e Biophoto Associates/Science, Photo Library: ti180064 - Fotolia: p.335 I Cl Manin Shiclds/Alamy, r Cl Fl.PA/Alamy;
p.107 aU Cl Michael Abbey/Science, Photo Library: p.108 Cl Wlm van p.337 Cl J Gmkn/Alamy: p.345 Cl Science VU/B. John. Visuals Un-
Egmond.Nisuals Unlimited, Inc/Science Photo Library; p.114 1 Cl A Limited/Science, Photo Library; p.351 Cl Mary Evans Picture Library/
Barrington Brown/Science Photo Library, m Cl Science Photo Llbra,y, Alamy: p.359 Cl Sally and Richard GreenhilVAlamy; p.363 Cl Graphic
r Cl Science Photo Library; p.129 I Cl Ed Reschke/Peter Arnold/Getty Science/Alamy; p.366 I Cl Frank Greenaway/Dorling Kinderslry/ Getty
Images. r Cl Dr Keith Wheeler/Science Photo Llbra,y; p.130 Cl Gene Images, m Cl alexfiodorov - Fotolia, re Frank Greenaway/Dorling
Cox : p.131 both Cl Geno, Cox; p.1 32 e C,.,ne Cox; p. 133 e C,.,ne Kindersley/C,.,tty Images: p.370 e Friedrich Siark/Alamy: p.380 I Cl
Cox; p.136 both e C,.,ne Cox, p.139 e C,.,ne Cox, p.141 1 Cl Bio- Geno, Cox, m Cl tina7si - Fotolia, r e Martin Fowler/Alamy; p.385 I Cl
photoAssociates/ScienccPhotolibrary,b Cl RichardJohnson; p.144 MindcnPicrures/SuperS!ock,r C ArlrianSherratt/Alamr; p.388 Clgre-
/e CJ Clegg, r Cl Gene Cox; p.147 e Biophoto Associa!es/Photo enwalcs/Alamy; p.392 both e CJ Clegg; p.394 I Cl Dave Wans/Alamy,
Researchers. lnc./Science Photo Library; p.153 Cl Gene Cox; p.155 m Cl Jon Durran!/Alamy, r Cl Nigel CauLin/Alamy; p.395 e wolfavni -
Cl Phototake lnc./Alamy: p.1 64 Cl Gene Cox; p.169 Cl doc-stock/ Fotolia; p.398 I Cl Craig Aumess/Corbi.s, m Cl inga spence/ Alamy,
Alamy: p.174 e S!e-vo, Gschmeissner/Science Photo Libra,y; p.1 75 t r Cl Tanguy Lo,Neel/Alamy; p.406 Cl tbkmedia.de/Alamy: p.407 l e
Cl Dr Gladden Willis, Visuals Unlimited/Science Photo Llbra,y, b Cl Danita Delimont/Alamy. r Cl All Canada Photos/Alamy; p.410 I Cl
BiophotoA.ssociates/SciencePhotoLlbra,y; p.l 761 Cl ScienceVU/ crisod - FotoLia,r Cl Suzannelong!Alamy; p.413 Cl PictureHooked/
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Alamy: p .177 ICI Andrzej Tokarski - Fotolia; p.179 ICI Phototake Inc./ p.418 I Cl Paul Heinrich/Alamy, re lsarpix/Alamy: p.422 Cl Manyn
Alamy: p.180 e CMEABG/UCB11. ISM/Science, Photo Library: p.182 1 F. Oi.illmaid/Science Photo Library; p.423 Cl Pau l Glendcll/ Alamy;
Cl Science, Photo Llbrary/ Alamy. r Cl Dr Tony Brain/ Science Photo p.428 CJ Clegg; p.429 1 Cl idp wildlife collection/Alamy, me Lip
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University Press, 1997); p .398Flgure17.20adap1edfromJulius
Janicke1. al.. PlamScience:Anlrnroduction!OWorldCrops.2nd
Past examioation questions reproduced by pennis.sion of University edition (W.H.Freeman, 1974); p.'!04 Figure 17.23 adapied from
of Cambridge lmernational Examinations A.D.Bradshaw and T. McNcilly. Evolution and PoUution (Cambridge
University Press. 1981),reproducedbypermissionofCambridge
An work and dam cr<.'tll!,o: University Press: p .44 1 Figure 18.24 IUCN Red List data (www.
pp.1'! 5.5 Table 7.3 adap!ed frcmj.Booner and A.W. Galston iucnredlisi.org), reproduced by permission of the lmemational Union
Principles ol Plam Physiology (W.H.Frceman, 1958); p.1 57 Figure for Conservation of Nature (IUCN): p .4S7 Figure 19.3 adapted from
8.6 from Colin Clegg (revised by S.lngham) from Exercise Physiology P. Moore. Recombioalll DNA Technology (The Biochemical Society,
and Functional Anatomy (Feltham Press, 1995), reproduced by 1994) Cl the Biochemical Society. reproduced by pennission of
permission of the author: p.1 66 Figure 8.15 H.G.Q. Rowen, adapied Portland Pres"' p.'! 66 Figure 19.11 from A.Mclenoan el.aL, Molecular
from BasicAna!omy & Physiology. 3/e (John Murray, 1987); p.184 Biology. 4!h Edition (Garland Science, 2013)
Table 9.2 from Raymond Pearl. 'fobacco Smoking and longevity'
from Science .. 2253 (4 Manch 1938); p.184 Table 9.3: Data on Permission for re-use of all Cl Crown Copyright information is
smoking, American Cancer Society, from http://txrwri!er.com/ gramed under the terms of the Open Government Licence (OGl).
onscience/Ospictures/smoltlngcancer2.2.jpg; p.185 Figure9.1S
fromCancerReseanchUK.httpJ/wv,,'W.cancerresearchuk.orsJ Thanksareduetothefollo"ingsourcesofinspirationanddatafor
canccr-infWcancerstats/type,;,llungs/indde nce; p.J88 Figure9.19 illustrative figures
World Health Organisation, from http://www.who.int/whr/1999/ p .59 Fig 3.3 Random collision possibilities adapied from Dianne Gull
eo/whr99_ch.S_en.pdf, Figure 9.20a National Institutes of Health, and Bernard Brown. 'Enzymes: fast and flexible'. Biological Sciences
from httpJ/en.wikipedia.orsfwiltl/File:Cancer_smoking_lung_ Rer:iew 6.2, November 1997; p.'!'!2 Fig 18.25 Adapied from Sruan 1
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World Health Organisa!ion, from wv,,w.who.int/whr/1999/eo/ Scientific Americ.:in Inc. 293:3
whr99_ch5_en.pfd(ISAugust,2007): p.1 94Figure 10.2re-usedfrom
Alas!ll.ir Gray, \lbrld Health and Disease (Open University Press Every effort has been made to trace all cop}Tight holders, but if any
198S). reproducedbypennissionofthe author; p . 197 Figure 10.7 have heenioadverten~yoverlooked the Publishers will he pleased to
World Health Organisation, from Weekly Epidemiological Record, make the necessary arrangements at the first opportunity.
71.(1996); p.202 Figure 10.14adap1edfromS.J .G. Kavanagh and
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